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ZooKeys 109: 1986 (2011)

The herpetofauna of Timor-Leste: a first report 19
doi: 10.3897/zookeys.109.1439 Research article
www.zookeys.org Launched to accelerate biodiversity research
The herpetofauna of Timor-Leste: a first report
Hinrich Kaiser1, Venancio Lopes Carvalho2, Jester Ceballos1, Paul Freed3, Scott
Heacox1, Barbara Lester3, Stephen J. Richards4, Colin R. Trainor5, Caitlin
Sanchez1, Mark OShea6
1 Department of Biology, Victor Valley College, 18422 Bear Valley Road, Victorville, California 92395, USA;
and The Foundation for Post-Conflict Development, 245 Park Avenue, 24th Floor, New York, New York
10167, USA 2 Universidade National Timor-Lorosae, Faculdade de Ciencias da Educaao, Departamentu da
Biologia, Avenida Cidade de Lisboa, Liceu Dr. Francisco Machado, Dili, Timor-Leste 3 14149 S. Butte Creek
Road, Scotts Mills, Oregon 97375, USA 4 Conservation International, PO Box 1024, Atherton, Queensland
4883, Australia; and Herpetology Department, South Australian Museum, North Terrace, Adelaide, South
Australia 5000, Australia 5 School of Environmental and Life Sciences, Charles Darwin University, Darwin,
Northern Territory 0909, Australia 6 West Midland Safari Park, Bewdley, Worcestershire DY12 1LF, United
Kingdom; and Australian Venom Research Unit, Department of Pharmacology, University of Melbourne, Vic-
toria 3010, Australia
Corresponding author: Hinrich Kaiser (hinrich.kaiser@vvc.edu)
Academic editor: Franco Andreone |Received 4 November 2010|Accepted 8 April 2011|Published 20 June2011
Citation: Kaiser H, Carvalho VL, Ceballos J, Freed P, Heacox S, Lester B, Richards SJ, Trainor CR, Sanchez C, OShea
M (2011) The herpetofauna of Timor-Leste: a first report. ZooKeys 109: 1986. doi: 10.3897/zookeys.109.1439
Abstract
Fieldwork conducted throughout Timor-Leste in September 2004 and July 2009 resulted in a collection
or recording of 263 herpetological specimens (100 amphibians, 163 reptiles), comprising at least seven
species of frogs and toads, 20 species of lizards, seven species of snakes, two species of turtles, and one
species of crocodile. Among the amphibians, the most frequently encountered species were toads (Dut-
taphrynus melanostictus), rice paddy frogs (genus Fejervarya), and rhacophorid treefrogs (Polypedates cf.
leucomystax). All three variants of rice paddy frogs encountered represent undescribed species similar to F.
verruculosa from neighboring Wetar Island. Records of F. cancrivora and F. limnocharis for Timor Island
are apparently errors based on misidentification. We obtained voucher specimens for a total of 147 lizards
and voucher photographs only for four specimens of Varanus timorensis. Aside from geckos frequently
associated with human habitations (e.g., Gehyra mutilata, Gekko gecko, Hemidactylus frenatus, H. platyu-
rus), we discovered an as yet undescribed species of bent-toed gecko, genus Cyrtodactylus, in the Same
valley. Our specimens of H. platyurus are the first record of this species from Timor-Leste. Commonly
encountered skinks included four-fingered skinks (genus Carlia), wedge skinks (genus Sphenomorphus),
and night skinks (genus Eremiascincus). Notable among the 15 snakes collected was the frequency of pitvi-
pers (Cryptelytrops insularis), which amounted to over 25% of all snakes. Our specimen of the wolfsnake
Copyright Hinrich Kaiser et al.. This is an open access article distributed under the terms of the Creative Commons Attribution License, which
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20 Hinrich Kaiser et al. / ZooKeys 109: 1986 (2011)
Lycodon subcinctus is the first record of this species for Timor-Leste. Based on these findings, it appears that
the biodiversity of amphibians and reptiles in this remote corner of Wallacea is much greater than previ-
ously thought, particularly with respect to scincid lizards. The detail we provide in the species accounts
is designed to allow the use of this report as a preliminary field guide to the amphibians and reptiles of
Timor-Leste. However, survey work is ongoing.
Keywords
herpetofauna, biodiversity, Timor-Leste, Wallacea
Introduction
Timor-Leste, also known as East Timor or Timor Lorosae, became the worlds newest
country on 20 May 2002 when its independence was restored after 24 years of Indonesian
occupation and three years of United Nations Transitional Administration. Prior to inde-
pendence in 1975, the country had nominally been a Portuguese colony since 1702, after
a continuous, though not unchallenged, Portuguese presence dating back to 1515. Its
remote location in the farthest southeastern reaches of Wallacea, the biogeographic region
delimited by Wallaces Line to the west, Lydekkers Line in the east, and the Timor Sea
in the south (fide Bellwood 2007), posed logistical challenges to all but the most widely-
traveled of biologists. The political uncertainty even during colonial times, particularly
in the eastern part of the island, was compounded by guerrilla and Indonesian military
activity over most of the past three decades and certainly proved a significant deterrent to
biological research. The countrys rugged terrain and an infrastructure that still reels from
the destruction wrought by the retreating occupation forces continue to make research
a challenge. Thus, herpetological survey work in Timor-Leste is still in its infancy. As a
consequence of the historical turmoil, there exist among the many dozens of specimens
from Timor Island in major herpetological collections around the world very few from the
eastern half of the island, the current territory of Timor-Leste. Here we present the results
from Phase I of the first comprehensive survey of the amphibians and reptiles in inde-
pendent Timor-Leste, along with an overview of the literature pertaining to the herpeto-
fauna of the country and a glossary to improve the use of this paper by non-specialists.
Geography and Geology. Timor Island (Portuguese: Ilha de Timor; Bahasa Indo-
nesia: Pulau Timor) is part of the Lesser Sunda Archipelago, an assemblage of islands
composed of a northern, volcanogenic arc (the Inner Banda Arc, with main islands Bali,
Lombok, Sumbawa, Flores, and Wetar) and a southern, orogenic arc (the Outer Banda
Arc, with main islands Sumba, Roti, and Timor). These islands are located in an area of
southeastern Wallacea (Fig. 1) where several biogeographic provinces converge. Timor
Island is the largest landmass in the area (30,777 km2), and it emerged as a landmass dur-
ing the Early Pliocene (ca. 5 mya) with a complex geologic history at the convergence of
the Australian continental plate and the Eurasian continental landmass (ESDD 2003).
The island is composed of a highly diverse stratigraphy, including several distinctive,
monolithic limestone karst formations (e.g., the Paitxau Mountains). By its position as
an oceanic landmass at the crossroads of the Southeast Asian and Australo-Papuan faunal
realms, the Timor Island herpetofauna could be expected to comprise a mosaic of diver-
sity created from these two main elements, with the addition of any endemic species.
The Democratic Republic of Timor-Leste occupies approximately the eastern half
of Timor island (15,410 km2), with the inclusion of the Oecusse-Ambeno coastal ex-
clave (Oecusse District; separated by an aerial distance of nearly 60 km west of the
nearest point in Bobonaro District, Timor-Leste), Jaco Island at the easternmost tip
of Timor (Lautm District; separated by a distance of less than 1 km from Tutuala
Beach), and Ataro Island (Dili District), 26 km north of the capital Dili. Whereas
Ataro (land area: 150 km2; Monk et al. 1997) is politically a part of Dili District in
Timor-Leste, it is volcanogenic in origin (age: 33.5 mya), geographically part of the
northern Banda Arc, and geologically distinct from Timor.
Habitats. Timor-Leste can be roughly divided into five major vegetation zones
(Monk et al. 1997), including thorn forest (dry coastal areas, primarily along the north
coast), dry deciduous forest (lower altitude habitats up to ca. 500 m), moist deciduous
forest, semi-evergreen rainforest (especially on slopes), and evergreen rainforest (in the
few pristine montane areas above 1000 m elevation). Trainor et al. (2007) provided a
more detailed account of habitat types, which we follow here. Their classification in-
cludes tall evergreen forest (tree height up to 40 m), semi-deciduous and tropical dry
forest types (tree height up to 20 m), a patchy tropical montane forest (elevations > 1000
m), beach forest and coastal scrub, savanna woodland, open eucalyptus forest, shaded
coffee plantations (> 600 m), swamps and swamp forests, rice paddies, and village land.
Habitats are generally characterized by sloping terrain (44% of the land in Timor-Leste
has a slope of 40%), rendering them unsuitable for sustainable agriculture (UNDP
2010). Whereas Timor-Leste is typical of the tropics in possessing only a thin soil layer,
there is little bare soil or grassland, and the island appears relatively well wooded.
There is little doubt that Timor-Leste was more forested before the arrival of the
Portuguese colonists in the early 16th Century, but it is also apparent that some types
of agriculture (such as the establishment of rice paddies) caused habitat modifications.
However, as first the colonial power and then the Indonesian occupiers exploited tropi-
cal woods (notably sandalwood and teak), the effects of ongoing shifting subsistence
agriculture became compounded. The reduction in the number of trees has by now
dramatically increased the threat of erosion during the infrequent but often torren-
tial rainfall, which may have serious consequences for road infrastructure. The threat
of continued deforestation to support unsustainable agriculture techniques and the
search for cooking fuel are real in Timor-Leste. These types of threats and the new
threat of invasive species make sustainability efforts imperative.
A brief history of herpetology in Timor-Leste. Exploration of the Lesser Sunda Ar-
chipelago began in earnest shortly after the explorers Louis-Antoine de Bougainville (in
1768) and James Cook (in 1770) sailed past the island (see van Aken 2005 for a review).
At this time, the well-established Dutch port of Kupang at the western end of Timor
(Coupang or Coepang in writings of the time) appears to have been a favorite place
for refitting on the globe-spanning voyages of the English commander Matthew Flinders
(expedition dates: 18011803) and the French commanders Nicolas Baudin (180004),
Louis-Claude de Freycinet (181720), Louis Isidore Duperrey (182225), and Dumont
dUrville (182629), all of whose vessels included naturalists and artists. It is remarkable
that any results at all were eventually reported from these voyages, since many of the
travelers died from diseases shortly after reaching Southeast Asia (among them the very
active Dutch collectors Heinrich Christian Macklot, Heinrich Boie, and Heinrich Kuhl).
The years from 18001830 may therefore be considered the first wave of scientific
exploration to the shores of Timor. During this initial wave, the most significant col-
lections in general, and on Timor in particular, were made by the French (e.g., Fran-
ois Pron and Charles-Alexandre Lesueur traveling with Baudin, Jean Ren Constant
Quoy and Joseph Paul Gaimard traveling first with Duperrey and then dUrville).
Whereas Baudin landed only in Kupang (in 1803), Duperrey visited both Kupang and
Dili (Diely or Dielly in writings of the time) in 1818. Shortly thereafter (1828),
the Dutch vessel Triton landed in Kupang with naturalists Macklot, Boie, and Salomon
Mller, who spent several years in the Lesser Sundas.
Even though in the late 1850s some amphibians and reptiles from Timor were sent
to the Dutch Rijksmuseum van Natuurlijke Historie (RMNH) by Pieter Bleeker, a med-
ical officer stationed in the Dutch East Indies, the second wave of exploration began in
earnest only with the explorations of the Swiss zoologists Paul Benedict Sarasin and Fritz
Sarasin, who traveled to the Lesser Sundas in 189396 and 190203, and continued
with the Siboga Expedition (18981900) under the leadership of Max Weber (RMNH).
This period also saw the publication of the only substantial Portuguese herpetological
reports on Timor (Bethencourt Ferreira 1897, 1898), the collections of Thomas Bar-
bour (190607; Museum of Comparative Zoology, Cambridge, Massachusetts, USA),
the extended museum-based reports by Nelly de Rooij (de Rooij 1915, 1917), as well
as the collections of Felix Kopstein (192224; RMNH), Malcolm Smith (1924, British
Museum of Natural History, London, United Kingdom), Emmett Reid Dunn (1926;
American Museum of Natural History, New York, USA), and Robert Mertens (1927;
Forschungsinstitut und Naturmuseum Senckenberg, Frankfurt am Main, Germany). Af-
ter this second wave had passed, a period of political instability commenced that included
the warfare of the 1940s, the period of decolonization during the 1960s and 1970s, and
the subsequent annexation by Indonesia, which halted research on Timor almost entirely
(with the exception of a collection made in West Timor by researchers from the Western
Australian Museum in the early 1990s) until Timor-Leste reached a modicum of politi-
cal stability within a few years of the 1999 UN-sponsored referendum on independence.
Timor Island and the southeastern corner of Wallacea are a particularly interesting
locale from a biogeographic point of view, but the relative influence of Southeast Asian
and Australo-Papuan elements on the herpetofauna has not been studied. Positioned
on the rugged eastern half of the largest landmass in the area (Timor is the 44th larg-
est island in the world and the 7th largest between Southeast Asia and New Guinea),
Timor-Leste is a likely source of significant endemism and well-positioned to begin an
assessment of the biogeographic provenance of its herpetofauna. In recognition of this
need, and given a near absence of herpetofaunal information for the country, we re-
solved to begin a comprehensive survey. This report summarizes the results of a survey
in 2004 (by SJR) and Phase I of a series of ongoing surveys conducted by the other
authors (except CRT and SJR) in July 2009.
Materials and methods

We collected amphibians and reptiles in Timor-Leste from 212 September 2004 (SJR
only) and from 1530 July 2009 at 18 localities throughout the country (Table 1; Fig.1).
Even though the country currently has not acceded to the CITES treaty, we decided a
priori to document the presence of known monitor lizards, pythons, and snake-necked
turtles primarily via photographic vouchers or from road-kills because of their CITES
Appendix II status, unless extenuating circumstances required adjustment of this proce-
dure. Were the species status of CITES-listed taxa in question, we would only collect a
minimal number of specimens and/or their tissues to ascertain their taxonomic status.
All other amphibians and reptiles were sampled and a sub-set of specimens encountered
was preserved as vouchers. During our survey we encountered some animals that were
not readily identified to species despite careful comparisons with type material and ex-
amination of pertinent taxonomic literature. In cases where there seemed to be superficial
resemblance to described taxa but where comparisons were complicated by the unavail-
ability or age of comparative material, by incongruities with our own field experience,
or by pending taxonomic revisions, we flagged the situation by inserting the clause cf.
(Latin: confer = compare) into the species name (e.g., Polypedates cf. leucomystax). Speci-
mens not matching any known species and presumed to be species new to science (can-
didate species, fide Padial et al. 2010) are first listed using the abbreviation sp. followed
by an integer and a candidate designation with a specimen number. For example, an un-
described species of Cyrtodactylus would be listed as Cyrtodactylus sp. 1 [Ca CMD 383],
where CMD 383 is the field number of a key specimen. Subsequent mentions of this
candidate species in the text will use only the abbreviation sp. with its assigned integer.
Localities. Whereas our surveys were conducted entirely within the country of
Timor-Leste, historical collections were concentrated in the western part of the island
near the historic port of Kupang. To avoid confusion, we refer to Timor when consider-
ing the entire island, but we may differentiate West Timor, defined as the western por-
tion of Timor and politically part of the Indonesian province East Nusa Tenggara (Ba-
hasa Indonesia: Nusa Tenggara Timur), from Timor-Leste (the sovereign nation). Timor
and Timor-Leste should not be confused, as has sometimes happened, with Timor Laut,
a name used for the Tanimbar Islands of Indonesias Maluku province to the northeast.
We conducted our surveys at localities (Table 1) that cover the major habitat types
and the diverse geography of the country (Fig. 2). Choice of localities was nearly always
influenced by the presence of supporting infrastructure (i.e., vehicle access, accommo-
dation, electricity) to facilitate collecting and processing of specimens. Since Timor-
Leste regained independence in 2002 (after a 7-day period in late 1975 just prior to
Indonesian annexation), efforts have been underway to reconstruct and solidify infra-
Figure 1. Map of districts, major towns, and collecting localities in Timor-Leste. Details for numbered
localities are listed in Table 1. Map by Mark OShea.
structure, such as roads, electricity, and telephone networks, but much of the existing
infrastructure remains in poor condition. This is especially the case the further one
travels from the capital, Dili.
Collecting. Survey protocols involved active searches along roadsides, forest paths,
and in riverbeds, supplemented by collection of road-kills and specimens encountered
fortuitously while driving. Searches were conducted during the day and at night. Dur-
ing the day we scanned the ground, tree trunks, and foliage, capturing frogs, lizards,
and non-venomous snakes by hand. We used neoprene-padded M1 snake tongs and
a Pro Bagger (Midwest Tongs, Independence, Missouri, USA) to handle, capture,
and transport venomous snakes. Lizards high in trees were stunned with plastic plugs
shot from 6-foot (183 cm) blowguns (Blowguns Northwest, Richland, Washington,
USA). We also removed bark from rotting logs and turned them over with stump
rippers (Midwest Tongs), carefully dismantled rock piles and similar potential refugia
by hand, shone flashlights into nooks and crannies, and raked through leaf litter. All
logs and rocks were replaced as closely as possible to their original position. By night
we used hand-held flashlights to search the ground, along the edges of watercourses
and swamps, and in the vegetation. As our collecting activities became known to the
local population, we were occasionally presented with specimens deemed worthy of
our attention (one lizard, two snakes, two turtles). These specimens were offered and
accepted with no request for remuneration. At the time of capture, we recorded date,
time, altitude, circumstances of capture, and GPS coordinates for each locality using a
Table 1. Localities surveyed in Timor-Leste during September 2004 and July 2009, including GPS
coordinates.
Locality District Locality Elevation (m) GPS1

Number
1 Dili area of Dili town and surrounds 120 S 0833' E 12532'
2 Ermera Eraulo village (Sta. Bakhita Mission) 1200 S 0847' E 12527'
3 Ainaro Eralisau village, near Maubisse 9011526 S 0850' E 12535'
4 Ainaro area of Maubisse town and surrounds 1484 S 0851' E 12536'
5 Ainaro slopes of Mount Ramelau 8312960 S 0852' E 12530'
6 Manufahi Turiscailau village, near Maubisse 1225 S 0850' E 12544'
7 Manufahi area of Same town and surrounds 513554 S 0900' E 12539'
8 Covalima area of Suai town and surrounds 13 S 0919' E 12515'
9 Viqueque Timor Village Hotel and surrounds 285 S 0847' E 12623'
10 Baucau area of Baucau town and surrounds 5350 S 0828' E 12628'
11 Lautem bat cave, Iliomar subdistrict 285 S 0839' E 12650'
12 Lautem area of Iliomar town and surrounds 315 S 0843' E 12650'
13 Lautem 40 min S Iliomar by road not determined S 0845' E 12649'
14 Lautem area of Lore 1 village and surrounds 3 S 0841' E 12659'
15 Lautem area of Lospalos town and surrounds 340 S 0831' E 12700'
16 Lautem 5 km S Mehara village 125 S 0827' E 12710'
17 Lautem Tutuala beach (Pantai Walu) 4 S 0825' E 12717'
1
GPS coordinates are approximate to define the area in which the survey work was carried out. Exact
localities are not provided to protect some of the unique and fragile habitats in Timor-Leste.
Garmin Oregon 400t (Garmin International Inc., Olathe, Kansas, USA). In order to
convey the relative frequency of observations (Table 2), we classified species encounters
as abundant (many encounters throughout a locality), common (usually present at a
locality), infrequent (few individuals seen, or unpredictable), rare (seen once only), or
indeterminate (in cases where our field experience and knowledge of the local popula-
tion was insufficient to permit classification).
Processing. During all surveys, at least one individual of each captured species at
each locality was photographed. Whenever possible, we also photographed the speci-
men in situ before capture. After capture, specimens were set up on a set, in a 90-
cm Cubelite (Lastolite Ltd., Coalville, Leicestershire, United Kingdom) using habitat
materials, and photographed in several positions to capture morphological detail in
life, and to ensure a record of all possible color patterns in life. Due to the vagaries of
specimen preparation in the field, we were unable to connect all photographic vouch-
ers shown in the figures to specimen numbers. Where possible, specimen numbers are
included in the figure captions.
Specimens were euthanized by intracardiac injection with a 5% procaine solution
(Altig 1980) or by immersion in dilute chlorobutanol (frogs only), following stand-
ard animal care protocols (e.g., ASIH 2004; Animals for Research Act Canada, RRO
1990, Regulation 24). Liver tissue samples were removed from most voucher speci-
mens through small lateral incisions and preserved in 1.5-ml plastic screw-cap centri-
Figure 2. Examples of sampled habitats in Timor-Leste. A Highland habitat in the area of Maubisse,
Ainaro District (altitude at the level of the buildings ca. 1400 m). This habitat has experienced consider-
able deforestation, as evidenced by the presence of small forest patches in the low-lying areas and the ab-
sence of trees on the higher slopes of Mt. Ramelau, in this view. This deforestation apparently began only
in the early 1980s (Trainor et al. 2007). The area supports a very active coffee industry. The patchwork of
forest, river valleys, coffee plantations, and deforested slopes creates a mosaic environment that most likely
will cater exclusively to habitat generalist species B Tiered rice paddies south of Baucau, Baucau District.
The area surrounding Baucau is a classic rice-growing region at low to moderate altitudes (sea level500
m). In this type of habitat we commonly encountered rice paddy frogs as well as their snake predators, and
some of the perianthropic geckos. Many of these terraced rice paddies have been operational for hundreds
of years, and they are a disturbed habitat exposed to strict human-initiated seasonality (a wet growing
season with artificial vegetation and irrigation, a dry fallow season with dry soil and absence of any vegeta-
tion) C Road leading through a lowland tropical evergreen forest (Trainor et al. 2007) on the southeastern
coast near Lor, Lautm District (altitude near sea level). This area also supports coastal dry forest, tidal
forests including mangroves, and coastal grasslands. The distance of this area from major population cent-
ers and its relative inaccessibility may be the primary reasons for the presence of such a diverse set of intact
habitats D The Lake Ira Lalaro floodplain and surrounding area. Whereas the foreground of this image
shows the marshy edges of the Irasequiro River, the background shows the treeless expanse of the lakes
floodplain. Because this area is a highly seasonal water source and prone to flooding, it has experienced
very little development. Photos by Hinrich Kaiser.
Table 2. Checklist of the species of amphibians and reptiles currently verified for Timor Island. The list of synonyms comprises those scientific names that have
been applied to Timor populations and is not an exhaustive list of synonyms for the species concerned. The most commonly used authorities are abbreviated as Ba
= Barbour (1912), BF97 = Bethencourt Ferreira (1897), BF98 = Bethencourt Ferreira (1898), Bl = Bleeker (1860), dH = de Haas (1950), dR15 = de Rooij (1915),
dR17 = de Rooij (1917), F = Forcart (1953), IC = Iskandar and Coljin (2001), S = Smith (1927), vK = van Kampen (1923), vL = van Lidth de Jeude (1895), Victor
Valley College survey = VVC. We consider as verified the occurrence of species either by our collection or by the presence of voucher specimens in museum collec-
tions that we have seen. Species in boldface print are confirmed for Timor-Leste. Altitudes listed are for specimens we collected. Habitat types follow Trainor et al.
(2007) and are abbreviated as TEF = tall evergreen forest, TDF = tropical dry forest, TMF = tropical montane forest above 1000 m altitude, BFCS = beach forest and
coastal scrub, SW = savannah woodland, EF = eucalyptus forest, CP = coffee plantations, SW = swamp and swamp forest, RP = rice paddies, and VIL = village land.
Frequencies listed are of those species we found in Timor-Leste, defined as abundant (many encounters at specific locality), common (usually present at a specific
locality), infrequent (few individuals seen, or unpredictable), rare (seen once), or indeterminate (in cases where our field experience and the knowledge of the local
population were insufficient to permit classification).
Current Name Synonyms (Authority) Altitude (m) Habitat(s) Frequency

Frogs and Toads
Family Bufonidae
Duttaphrynus Duttaphrynus melanostictus (VVC) 0600 TDF, BFCS, SW, CP, abundant
melanostictus RP, VL
Bufo melanostictus (Trainor et al. 2009)
Family Dicroglossidae
Fejervarya spp. Rana tigrina (Bl) 01200 TEF, TMF, CP, SW, RP, abundant
VL
Rana tigerina var. verruculosa (Roux 1911)

Rana tigerina (Ba)
Rana verruculosa1 (S, vK, Menzies 1987)
Rana cancrivora (S, vK)
Limnonectes timorensis Limnonectes timorensis (VVC) > 1000 TMF infrequent
Rana timorensis (S)
Hylarana elberti (F, Menzies 1987)
27
28

Family Hylidae
Litoria everetti Litoria everetti (VVC) > 1000 TMF infrequent
Hyla everetti (Ba, S, vK),
Family Rhacophoridae
Polypedates cf. leucomystax Polypedates cf. leucomystax (VVC) 01400 TEF, TDF, TMF, SW, common
EF, CP, SW, VL
Polypedates leucomystax (Ba, Bl, vK)
Rhacophorus leucomystax (vK)
Rhacophorus leucomystax var. sexvirgata (vK)
Lizards
Family Agamidae
Draco timoriensis Draco timoriensis (VVC, Ba, Bl, dR15) 0300 TEF, TDF, BFCS, VL
Draco timorensis (BF98, vL, Manaas 1972),
Draco viridis var. timoriensis (Schlegel 183744)
Draco haematopogon (Bl)
Draco lineatus (Bl)
Draco volans (dR15)
Draco walkeri (Ba, dR15)
Family Gekkonidae
Hinrich Kaiser et al. / ZooKeys 109: 1986 (2011)
Cyrtodactylus sp. Cyrtodactylus sp. (VVC) 600 TEF, CP rare

Goniodactylus Timorensis2 (Boulenger 1885)
Gymnodactylus (?) marmoratus (S)
Gehyra cf. mutilata Gehyra cf. mutilata (VVC) Lowlands TDF, BFCS, SW, EF, infrequent
RP, VL
Gehyra mutilata (dR15)
Peropus mutilatus (S)
Hemidactylus platurus3 (Bl)
Gekko gecko Gekko gecko (Ba, VVC) 0300 TEF, TDF, BFCS, SW, common
EF, CP, VL
Gekko verticillatus (BF98, dR15, vL, Manaas 1972)
Platydactylus guttatus (Bl)
Hemidactylus frenatus Hemidactylus frenatus (BF98, Ba, Bl, dR15, VVC) 0300 TDF, BFCS, SW, EF, abundant
CP, RP, VL
Hemidactylus platyurus Cosymbotus platyurus (VVC) 0300 TDF, BFCS, SW, VL infrequent
Family Scincidae
Carlia spp. Carlia spp. (VVC) 01500 TDF, TMF, BFCS, VL common
Carlia peronii Carlia peronii (Zug 2010)
Heteropus peronii (Dumril and Bibron 1839)
Carlia spinauris Carlia spinauris (Zug 2010)
Lygosoma (Leiolopisma) spinauris (S)
Cryptoblepharus Cryptoblepharus leschenault (VVC) Lowlands TDF, BFCS infrequent
leschenault
Ablepharus boutonii leschenault (F, Mertens 1930)
Eremiascincus spp. Eremiascincus spp. (VVC) 02100 TDF, TMF, BFCS common
Eremiascincus antoniorum Glaphyromorphus antoniorum (Greer 1990)

Lygosoma (Omolepida) antoniorum (S)

Eremiascincus timorensis Glaphyromorphus timorensis (Greer 1990)
Eutropis cf. multifasciata Eutropis cf. multifasciata (VVC) 01200 TDF, SW, EF, CP, VL common
Mabuya multifasciata (dR15, S)
Euprepes Sebae (Bl)
29
30

Lamprolepis cf. Lamprolepis cf. smaragdina (VVC) 0300 TDF, BFCS, SW, EF, common
smaragdina VL
Lygosoma smaragdinum (Bl, dR15, vL)
Dasia smaragdinum (Ba)
Sphenomorphus spp. Sphenomorphus spp. (VVC) 01500 TEF, TDF, TMF, BFCS infrequent
Family Varanidae
Varanus timorensis Varanus timorensis (BF98, dR15, S, vL, Boulenger Lowlands BFCS, SW, RP, VL infrequent
1885, VVC)
Varanus timoriensis (Ba, Bl)
Snakes
Family Colubridae
Coelognathus subradiatus Coelognathus subradiatus (VVC) 0400 TDF, VL infrequent
Coluber melanurus4 (dR17)
Coluber melanurus var. timoriensis (BF97, BF98, dR17)
Coluber subradiatus (dR17, S)
Compsosoma melanurus4 (Bl)
Compsosoma subradiatum (Bl)
Elaphe melanura (Ba)
Elaphe subradiata (dH, Schulz 1996)
Elaphe timoriensis (Ba)

Elaphis subradiatus (vL)
Dendrelaphis inornatus Ahaetulla picta inornata5 (dH) 0300 TDF, BFCS infrequent
timorensis
Dendrelaphis inornatus timorensis (F), VVC)
Dendrophis picta (Bl)
Dendrophis pictus (Ba, BF98, dR17, vL)
Dendrophis pictus timorensis (S)
Lycodon capucinus Lycodon aulicum (Bl) 500 TDF, VL indeterminate
Lycodon aulicus (Ba, dR17, S)
Lycodon aulicus capucinus (dH)
Lycodon aulicus var. D (BF98, Boulenger 1893)
Lycodon capucinus (VVC)
Lycodon subcinctus Lycodon subcinctus (VVC) 500 VL indeterminate
Family Homalopsidae
Cerberus rynchops Cerberus rynchops (BF98, dR17, VVC) Lowlands RP infrequent
Cerberus rynchops rynchops (dH)
Cerberus boaeformis (Bl)
Hurria rhynchops (Ba)
Family Typhlopidae
Ramphotyphlops braminus Ophthalmidion crassum6 (Bl) 0300 TDF, VL infrequent
Ramphotyphlops braminus (VVC)
Typhlops braminus (Ba, dH, dR17, vL)
Family Viperidae
Cryptelytrops insularis Cryptelytrops insularis (VVC) 0500 TDF, BFCS, EF, RP, VL infrequent
Bothrops erythrurus (vL)
Bothrops viridis (Bl)
Lachesis gramineus (BF98, dR17, Manacas 1972)

Trimeresurus albolabris (dH)
Trimeresurus gramineus (Ba)
Turtles
Family Chelidae
Chelodina timorensis Chelodina timorensis (McCord et al. 2007) 300 Lake Ira Lalaro rare
Chelodina mccordi timorlestensis (Kuchling et al. 2007)
31
32

Chelodina novae-guineae (Ba)
Family Geoemydidae
Mauremys reevesii Mauremys reevesii (Kaiser et al. 2010) 0-300 VL rare
CroCodiles
Family Crocodylidae
Crocodylus porosus Crocodilus biporcatus (Bl) Lowlands SW common
Crocodilus porosus (BF98)
Crocodylus porosus (Ba, dR17)
1
Menzies (1987) recorded vocalizations of rice paddy frogs, which he called a short call (Menzies 1987: Fig. 16b) and a long call (Menzies 1987: Fig. 16c). We
have heard these types of calls and traced them to males of what we believe to be two distinct species of Fejervarya.
2
Grays (1845) and Boulengers (1885) Catalogues list this species in the synonymy of what is now known as Cnemaspis boiei, a gecko from India. Grays descrip-
tion of the specimen from Timor reads thus: In spirits. Wants tail. Timor ? Presented by T. Bell, Esq. The fact that this tailless specimen was from a different
collection than that received by Dumril and Bibron in Paris, which itself consists of only a single bent-toed gecko specimen from Timor, raises the possibility
that a species of Cyrtodactylus or Cnemaspis other than the one described by Dumril and Bibron (1837) was discovered by Thomas Bell among the material
collected on the voyages of HMS Beagle. Gray (1845) writes that the specimens procured by Darwin and Captain Fitzroy were presented to the British Museum
of Natural History by Bell after describing them in his volume titled Zoology of the HMS Beagle. The specimen listed by Gray (1845) is presumed lost (C. Mc-
Carthy, in litt.).
3
Bleeker (1859) provided a detailed listing of a form he named Hemidactylus platurus. This form is related to Gehyra mutilata based on the original description,
and it is therefore not to be confused with H. platyurus. However, since no type specimen was designated, the name H. platurus Bleeker (1859) is a nomen
nudum.
4
The names Coluber melanurus, Compsosoma melanurus, and Elaphe melanura are currently in the synonymy of Coelognathus flavolineatus. Absent any specimens
of C. flavolineatus from Timor and given its known range, we consider all reports of this species on Timor in error and refer them to C. timoriensis.
5
According to How et al. (1996), A. p. inornata is a synonym of D. i. timorensis.
6
Bleeker (1860) listed this name in error for what is undoubtedly a reference to R. braminus. The name Ophthalmidion crassum is actually a synonym of the South
American blindsnake Typhlops reticulatus.
fuge tubes containing 95% non-denatured ethanol (EtOH). Ancillary collections of

external and internal parasites (mites, ticks, tapeworms, nematodes) were made op-
portunistically when such organisms were discovered during specimen processing. In
instances where captive individuals defecated into their plastic bag, fecal samples of the
fresh material were preserved in 2.5% aqueous (w/v) potassium dichromate (K2Cr2O7)
according to the method described by Duszynski et al. (1982). Immediately after eu-
thanasia, snout-vent length (SVL) and total length (TL) were measured for all reptiles
to ensure that specimens could be laid out straight and measured accurately. We also
took some scale counts to aid in the identification of species. Specimens were sexed
before preservation when possible by everting hemipenes or by checking for the pres-
ence of ovaries or yolked eggs. For lizards, data on maturity and reproductive condition
were made after preservation and will be reported elsewhere (Goldberg et al., in prep.).
All specimens were fixed in 10% formalin. Amphibians were placed directly into
plastic tubs and carefully positioned, whereas reptiles were first injected with 10% for-
malin and then positioned. Specimens were then covered with formalin-saturated pa-
per towels before the tubs were sealed and the specimens left to set. After fixing, field-
numbered series of potentially problematic taxa (e.g., gekkonids, scincids, anurans)
were photographed in dorsal and ventral view for later comparative study. Voucher
specimens of amphibians and reptiles (Appendix I) have been deposited in the Divi-
sion of Amphibians and Reptiles, National Museum of Natural History, Smithsonian
Institution, Washington DC, USA (USNM) and the South Australian Museum, Ad-
elaide, Australia (SAMA). At the time of publication, some specimens are listed for the
USNM using CMD field numbers, as these have not yet been accessioned. However,
these specimens are scheduled to become permanently integrated into the USNM
collection by August 2011, at which time they will be accessible for research. Further-
more, searches of the USNM specimen database using these field numbers will reliably
yield the specimens designated by these numbers herein.
Supplement. In addition to the information provided in this paper, a brief glos-
sary of relevant scientific terms is provided on the Zookeys website to augment public
understanding of the information presented herein, and to allow local authorities the
use of this paper as an instrument for recognizing the herpetofauna of Timor-Leste.
Species accounts
In the following accounts, we provide the most current accepted scientific name for
each species based on literature available as of 1 September 2010. Each species name
is given with its author to make its taxonomy unequivocal. A more complete checklist
of species known from Timor and Timor-Leste with synonyms used to document their
presence is provided in Table 2. Beneath the species name we list common names in
English (E), taken from or modeled according to common usage by professional her-
petologists, and in Tetun (T) and any local language for which a name is commonly in
use. Tetun names with an asterisk (*T) are newly coined and designed to approximate
the English name. Words in Tetun are spelled based on Hull (2001), with localities
spelled according to common usage and in recognition of historic names (C. Williams-
van Klinken, in litt. 20 July 2010). Our accounts include brief descriptive statements
to aid field identification, taxonomic comments, information regarding the natural
history of species, and any additional comments that may assist the readers under-
standing of the species occurrence in Timor-Leste. The list of specimens examined is
provided in Appendix I in the same order as species are listed in the main text.
Frogs and Toads (Order Anura)

Family Bufonidae True Toads
Duttaphrynus melanostictus (Schneider, 1799)
http://species-id.net/wiki/Duttaphrynus_melanostictus
Fig. 3
Common names. (E) Black-spined Toad, Common Asian Toad, Common Sunda
Toad. (T) Manduku Interfet (manduku = frog, INTERFET = International Force for
East Timor; see below).
Identification. This toad can be recognized by its stout body, dry and warty skin,
and by the distinct pattern of bony ridges (cranial crests) on the head. The shape and
color of these ridges are characters useful for distinguishing among different toad spe-
cies. In D. melanostictus they are of varying thickness and include a canthal ridge,
supralabial ridge, and a series of ridges framing the eye (preorbital, supraorbital, pos-
torbital, and orbito-tympanic ridges; Fig. 3). The tops of these ridges are usually black.
A second important and useful characteristic to differentiate between toad species is
the size and shape of the large parotoid gland (sometimes also described as a poison
gland) on either side of the head. In D. melanostictus this gland is elongate and about
23 times the size of the eye (Fig. 3). This species of toad also usually has several pro-
truding wart-like skin glands on its back behind its head in addition to many smaller
glands all over its body, most of which are tipped with black. Our familiarity with the
species from elsewhere in the region allows us to confirm its identity.
Collection and natural history. Hiding under a diverse array of objects by day
and active by night, this toad is a relatively recent invader of Timor-Leste (see Trainor
2009), yet it was one of the most commonly encountered amphibians where it oc-
curred. The distribution of D. melanostictus in Timor-Leste currently excludes areas
of high elevation (above ca. 1200 m) as well as the region east of a line connecting
Manatuto and Viqueque (Trainor 2009). We collected five voucher specimens at night
(e.g., on the path leading to the Trilolo River north of Same, Manufahi District, alti-
tude 553 m) and noted the presence of this species in disturbed habitats (e.g., towns,
roadsides), cultivated habitats (e.g., coffee plantations) and some fairly pristine habitats
(e.g., coastal scrub). Individuals ranged from tadpoles and juveniles (though not re-
cent metamorphs) to adults and they exhibited varying shades of dull yellow to brown
Figure 3. Duttaphrynus melanostictus. Yellow morph from Same, Manufahi District (SVL 53 mm). Photo
by Mark OShea.
coloration. This is the only species of true toad reported from Timor-Leste so far, but
we have been unable to verify the presence of specimens from Timor in herpetological
collections. Therefore, our records appear to be the first vouchered confirmation of this
species for Timor-Leste and Timor.
Toad introductions. Reports by Australian peacekeepers of the cane toad, Rh-
inella marina, in Timor-Leste are an error arising from the soldiers familiarity with
R. marina, the only bufonid introduced to Australia and New Guinea, and their
lack of familiarity with the Asian D. melanostictus. It is interesting in this regard
that this species has taken on the Tetun name of the International Force for East
Timor (INTERFET), the transitional peacekeeping force that arrived to stabilize
the country after the departure of the Indonesian occupation force. INTERFET was
composed primarily of Australian peacekeepers and the local belief, based on the
erroneous identification of D. melanostictus by these personnel, is that INTERFET
is responsible for the presence of this toad in Timor-Leste. The distribution of D.
melanostictus includes several other Lesser Sunda Islands (e.g., Bali, Lombok), and it
may be impossible to ascertain from where and when the initial wave of toad inva-
sion originated.
The introduction of toads to non-native environments has frequently resulted in
ecological disasters. The most notorious example of this has been the cane toad (R.
marina), whose spread by humans has become a problem with nearly global impli-
cations (e.g., Covacevich and Archer 1975; Lever 2001; Phillips et al. 2007). The
species has been carried from its native northern South American habitat to locations
as widespread as the Greater and Lesser Antilles, Florida, Hawaii, the Fiji Islands,
the Philippines, Taiwan, the Ryukyu Island Archipelago of Japan, several Pacific is-
lands, New Guinea, and, famously, Australia (see Zug and Zug 1979). It appears
that the spread of D. melanostictus may rank a close second in terms of its geographic
reach (from the Indian subcontinent throughout mainland and insular Southeast
Asia), though perhaps not in terms of its ecological significance (e.g., Inger and
Voris 2001). However, toads are voracious opportunistic predators whose impact
on a newly colonized ecosystem may take years to assess. Reported impacts include
alteration of the food chain, detrimental effects on lizard population recruitment,
extirpation of leaf litter amphibians and their tadpoles, reduction of amphibiopha-
gous reptile and mammal densities, and even poisoning of human or canid preda-
tors (e.g., Trainor 2009). We have recommended to government agencies that the
advance of D. melanostictus in Timor-Leste requires close monitoring and a popular
campaign to avoid human injury. A second toad invasion appears to be underway
concurrently by Ingerophrynus biporcatus (formerly Bufo biporcatus) on Roti Island,
an island neighboring Timor (Trainor 2009).
A simple distinction between D. melanostictus and R. marina can be made by look-
ing at the morphology of features described above. Whereas D. melanostictus has an
elongated parotoid gland that is about three times the size of the eye (Fig. 3), the gland
of R. marina is considerably larger (nearly five times the size of the eye) and shaped
like an irregular rectangle with rounded corners. The cranial crests of D. melanostictus
are relatively thin and topped with black, whereas those of R. marina are rather stout,
surround the eye and are colored as the rest of the head.
Taxonomic comment. Prior to the revision of amphibian taxonomy by Frost et al.
(2006), this species was known as Bufo melanostictus.
Family Dicroglossidae Fork-tongued Frogs

Genus Fejervarya
http://species-id.net/wiki/Fejervarya
Fig. 4
Common names. (E) Rice Paddy Frogs. (T) Manduku natar (manduku = frog, natar
= rice paddy).
Identification. Rice paddy frogs (Fig. 4) are the most common amphibians found
in regions with rice agriculture. They may grow to over 60 mm in snout-vent length
and can usually be recognized by their fairly stout body shape, brownish to gray-green
coloration, shiny moist skin with ill-defined dorsal and lateral patterns, and warts scat-
tered irregularly or in rows along the back (e.g., Fig. 4B).
Collection and natural history. As the name suggests, rice paddy frogs are com-
monly found in rice paddies where they perch at the waters edge, on tufts of vegeta-
tion, or even on cow patties. Our survey documented at least three species of rice
paddy frogs in Timor-Leste, with two or more often occurring in the same suitable
habitat. Species can be distinguished by the size of mature males, which can be clearly
separated into three groups by their size, morphology of the tympanum and its as-
sociated structures, and the patterning of the throat in males. One of the species,
designated as Fejervarya sp. 1 [Ca CMD 431] is a lowland form and most similar to F.
verruculosa (Roux 1910). This candidate species differs from F. verruculosa by the shape
of the supratympanic fold, the size of the tympanum relative to the eye, patterning on
the hidden surfaces of the legs, and number and position of maxillary teeth and the
shape of the alary process of the premaxilla, among other characters. The other two
candidate species can also be differentiated by these and other characters, and they have
been designated Fejervarya sp. 2 [Ca CMD 508] and Fejervarya sp. 3 [Ca CMD 355].
Among the specific habitats where we encountered these frogs were active rice
paddies, roadside puddles, coffee plantations, and coastal forests at altitudes between
4 m and 1187 m. Our observations are consistent with those of Menzies (1987) but
we believe that not all three presumptive species are capable of such habitat plasticity.
Figure 4. Rice paddy frogs, genus Fejervarya. A Fejervarya sp. 1 from the Baucau area (SVL 58 mm) BFe-
jervarya sp. 2 from the Viqueque area (SVL 46 mm) C Fejervarya sp. 3 from the Viqueque area (SVL 38 mm)
DAll three species of rice paddy frogs found in Timor-Leste may show varying degrees of green coloration on
the dorsal and lateral parts of the body. This specimen (SVL 56 mm) from the Viqueque area represents the
most extreme green coloration we observed, in terms of both brightness and coverage. Photos by Mark OShea.
More detailed investigation is needed to clarify the habitat requirements of these three
candidate Fejervarya species.
Biogeography. The presence in Timor-Leste of three morphologically similar and
seemingly endemic dicroglossid frogs, with an evolutionary origin in Asia, raises inter-
esting biogeographic questions. The simplest explanation would be a single or a series
of introductions in modern times, with the influx of peoples and cargo from points all
across the Indonesian Archipelago. However, a more ancient, classic island biogeogra-
phy scenario is also feasible. Based on the two main concepts of speciation, sympatric
speciation among amphibians is possible but presumed rare among dicroglossid frogs.
The concept of allopatric speciation is the alternative, by which the three species may
represent descendants of three separate introductions that occurred as early farmers
brought rice plants to the island. It is generally accepted that rice agriculture origi-
nally spread from China into South and Southeast Asia (Crawford and Chen 1998)
and reached the Lesser Sunda Islands in waves after spreading throughout the Greater
Sunda Islands during the Neolithic Period (Chi and Hung 2008). During this time, it
is quite possible that stowaway frogs arrived on Timor with rice plants. The occurrence
of several similar rice paddy frog species in sympatry is not unique to Timor-Leste
(e.g., Burma; G. Zug, in litt.). Molecular studies to obtain some insights into this
conundrum are progressing. The hypothesis that multiple human-mitigated introduc-
tions of Fejervarya populations occurred mirroring the development of rice cultivation
is a plausible explanation for the many species of this genus listed as incertae sedis with
respect to their intrageneric relationships.
Limnonectes timorensis (Smith, 1927)

http://species-id.net/wiki/Limnonectes_timorensis
Fig. 5
Common names. (E) Timor River Frog. *(T) Manduku mota (manduku = frog, mota
= river).
Identification. Frogs of the genus Limnonectes are nocturnal and generally quite
difficult to distinguish from similar species. Limnonectes timorensis is currently the
only known species of river frog recorded from Timor-Leste. Identifying characteris-
tics include fingertips that are slightly swollen and widened at their tips, but which
do not possess a marginal fold that outlines the disk pad. The first finger is invariably
longer than the second. They possess a dorsolateral fold that originates just behind
the eye and continues dorsally to the groin, and a tympanum that is nearly equal to
the size of the eye (Fig. 5). A brown band is present on the head, arising near the tip
of the snout, continuing along the canthus rostralis through the eye, and completely
enveloping the tympanum. The skin warts commonly found concentrated on the
dorsum in other species of Limnonectes are reduced in number on the dorsum but
quite prominent on the side of the body (Fig. 5). We were readily able to confirm our
Figure 5. Limnonectes timorensis. Female from near Eraulo, Ermera District (USNM [CMD 422], SVL
62 mm). Photo by Mark OShea.
identification by consulting the figure presented in Smith (1927: Pl. II, Fig. 1) and
from the original description.
Variation. Whereas in one of our specimens the internarial distance is slightly
greater than the interorbital distance, a diagnostic characteristic provided by Smith
(1927), the internarial distance is equal to the interorbital distance in the second.
Even though Smith (1927) did outline some variation among his eight specimens,
only slight differences in the interorbital distance are mentioned. It appears that the
comparison of internarial distance with interorbital distance by itself is insufficient to
distinguish this species from others.
In a second instance of incongruity between our specimens and the original de-
scription, the nares are not at the midway point between the eye and the tipof the
snout but located approximately one third of the eye-to-snout distanceaway from the
snout in both specimens. We believe that this incongruity could be due to an error in
Smiths (1927) description sincethe drawing of the frog (Smith 1927: Plate II, Figure
1) conforms to our specimens and not to Smiths description. Scientific illustrators are
generally extremely meticulous and accurate, and misplacement of the nares, a key
character of the head, would be an unlikely error.
Lastly, the foot in our specimens is 7% longer than, as opposed to equal to, tibia
length as described by Smith (1927). Repeated measurements with digital calipers and
mechanical calipers as used in Smiths day resulted in measurement errors of 3% and
7%, respectively (n = 10 for each instrument). We therefore believe that the discrepan-
cy between foot and tibia length measurements we made on our specimens, and Smith
made on his, is due to slight variation in the hindlimbs (Smith 1927:212), combined
with measurement error, and does not represent a diagnostic difference.
Collection and natural history. Two female individuals were collected during a
single night from the Meleotegi River, near Eraulo, Ermera District, altitude 1179 m.
During the dry season, the Meleotegi River is a relatively shallow stream that runs over
pebbles and allows easy crossing. Boulders are distributed at irregular intervals along
and in the riverbed. It is clear from the steeply eroded riverbanks (over 5 m high in
some parts) that the river carries a large volume of water during parts of the year. One
individual of L. timorensis was collected from a thin branch overhanging the relatively
steep riverbank, whereas the other was found on a large boulder in midstream. No
vocalizations were heard.
Taxonomic comment. Forcart (1953) and Menzies (1987) considered L. timoren-
sis a synonym of Hylarana elberti Roux 1911. Forcarts (1953) synonymy was based on
a comparison of specimens in the Naturhistorisches Museum Basel, Switzerland with
the holotype of H. elberti by Robert Mertens. One of us (HK) has made a very careful
comparison of our L. timorensis specimens with the holotype of H. elberti, and found
differences in the tuberculation of the hand, the width of fingertips, the position and
size of the rictal gland at the angle of the jaw, the extent of toe webbing, the width of
the toe tips, and the patterning and consistency of the skin on the throat. Additional
features of the shape of the head are difficult to compare because the holotype of H.
elberti has a damaged anterior of the head. We therefore agree with Dubois (1986) that
the species from Timor and Wetar are distinct.
Family Hylidae Treefrogs

Litoria everetti (Boulenger, 1897)
http://species-id.net/wiki/Litoria_everetti
Fig. 6
Common names. (E) Everetts Timor Treefrog. *(T) Manduku ai Timor (manduku =
frog, ai = tree).
Identification. Litoria everetti can easily be recognized by a combination of the
following traits: webbed hands and feet, expanded finger and toe tips or disks, a well-
developed supratympanic fold (Fig. 6). The hidden portions of the legs have an orange
marbled pattern in life. In common with most treefrogs, this species is nocturnal. Men-
zies (1987) provided an excellent description of this species and a photograph. SJR is
familiar with this species and could confirm its identity.
Figure 6. Litoria everetti. Female from near Eraulo, Ermera District (USNM [CMD 420], SVL 59 mm).
Photo by Mark OShea.
Collection and natural history. Two females of L. everetti were collected at night
at the same locality and during the same night as Limnonectes timorensis. Based on
their overall morphology and characteristic webbing, as well as their characteristically
orange marbled thighs (Boulenger 1897), these are clearly L. everetti. One specimen
was found at the edge of the riverine embankment, perched on a branch. The second
was caught on a small boulder near the waters edge. Litoria everetti is a species in Red
List Category Least Concern (Iskandar and Mumpuni 2004).
Family Rhacophoridae Afro-Asian Foam-nest Treefrogs

Polypedates cf. leucomystax (Gravenhorst, 1829)
http://species-id.net/wiki/Polypedates_leucomystax
Fig. 7
Common names. (E) Striped Treefrog, Four-lined Treefrog, Golden Treefrog. (T)
Manduku ai-riskadu (manduku = frog, ai = tree, riskadu = striped) or manduku loron
(manduku = frog, loron = sunlight). (Fataluku) Nelu cila. The common names in Te-
tun and Fataluku are generalized name for treefrogs and may be applied to other such
species without detailed distinction.
Identification. This species is a relatively slender treefrog with a variety of dor-
sal patterns (Fig. 7). The background coloration is usually a light brown during the
Figure 7. Polypedates cf. leucomystax. The individuals shown display the diversity of color patterns found
in this species A A specimen from Bakhita (SVL 45 mm) displaying irregular dark brown spots and barred
legs on a lighter brown background B A specimen from Lor, Lautm District (SVL 48 mm), presenting
with a combination of brown dorsal and dorsolateral lines and leg barring on a nearly yellow background
C A specimen (SVL 46 mm) from the same locality as B, showing a very lightly colored dorsum devoid
of lines and spots. Photos by Mark OShea.
nightly activity period but becomes a deeper brown while individuals are resting in
their diurnal refuges. Patterning may consist of darker lines or bands, brown blotches,
crossbars on limbs, or there may be no pronounced pattern. In comparison with the
two other potential tree-dwellers encountered, Limnonectes timorensis and Litoria ever-
etti, P. cf. leucomystax is more slightly built, has a pointed snout, lacks raised warts or
tubercles on the back, and has no webbing between the fingers.
Collection and natural history. These frogs were quite commonly found, call-
ing from the edges of ponds (e.g., 6.0 km W Lor 1 village, Lautm District), from
small shrubs and bushes (e.g., 5 km S Malahara, Lautm District), from the trunks of
fallen trees, and from boulders in the middle of streambeds (e.g., near Timor Village
Hotel, Wailakurini, Viqueque District), at altitudes from near sea level to 1350 m at
Maubisse, Ainaro District.
Taxonomic comment. Our experience with the geographically widespread rhaco-
phorid usually identified as Polypedates leucomystax, with a range extending from Bor-
neo to Peninsular Malaysia, and from India to Cambodia, is that it is a taxon in need
of closer investigation. On a recent trip we observed the species in Sabah, Borneo, and
shortly thereafter in Timor-Leste. Based on vocalizations, behavior, maximum size,
and color and pattern variation, we cannot confidently assign our specimens to P. leu-
comystax and instead refer to them as P. cf. leucomystax. The widespread morphotypes
collectively known as P. leucomystax are likely an assembly of similar rhacophorid spe-
cies that occupy a treefrog niche in geographically distinct locations, as others have
suggested (e.g., Narins et al. 1998). The taxonomic conundrum presented by P. leuco-
mystax is currently the subject of both molecular and morphological study (e.g., Brown
et al. 2010), but a resolution has so far been elusive.
Lizards (Order Lacertilia)

Family Agamidae Agamas and Dragons
Draco timoriensis Kuhl, 1820
http://species-id.net/wiki/Draco_timoriensis
Fig. 8
Common name. (E) Timor Flying Dragon, Timor Flying Lizard. (T) Teki liras (teki =
gecko, liras = winged). Fataluku: Lika. Mambae: Berdigil.
Identification. Lizards of the genus Draco are diurnal and easily identified by
the presence of patagia. These wing structures (Fig. 8) consist of skin flaps that are
stretched across highly modified ribs that allow the lizards to glide between trees. Al-
though referred to as flying lizards these and other reptiles that have perfected this
escape strategy are actually only gliding, flight being the preserve of birds, bats, and
insects. They also possess a dewlap under the chin that males use for territorial display.
Draco timoriensis is the only species of its genus known to occur on Timor (see taxo-
nomic comments below).
Figure 8. Draco timoriensis. Male from Wailakurini, Viqueque District (USNM 573658, SVL 75 mm,
TL 208 mm). Photo by Hinrich Kaiser.
Collection and natural history. We captured four specimens of D. timoriensis and

observed several others. All individuals were initially seen high off the ground (> 5 m)
on the trunks or larger branches of trees but never on palm trees. Even though they are
cryptically patterned against the bark background when stationary against the trunk of
the tree, they are easily spotted when displaying their bright yellow dewlaps (Fig.8).
Our specimens were captured either using blowguns or by climbing the tree and forc-
ing the lizard to glide to an accessible height. All specimens were seen and captured
during the daytime. Where they occurred, these lizards were not rare. However, their
dispersal pattern appears to be clumped (several lizards in one area with none outside
of a particular territory) and we did not discern any pattern to their localized distribu-
tion. Based on our encounters, D. timoriensis is limited in its distribution to altitudes
from sea level to ca. 300 m.
Taxonomic comments. Several historic reports of Draco collected on Timor list D.
volans or D. walkeri in addition to D. timoriensis. Based on recent unpublished findings
from a molecular analysis (J. McGuire, in litt. 13 Oct 2009), D. volans is confined to
Bali and Java whereas the distribution of D. walkeri is limited to Sulawesi (McGuire
et al. 2007). All records for flying lizards from Timor should therefore be attributed to
D. timoriensis.
The species name of the Timor flying lizard has variously been spelled timorien-
sis or timorensis. In the accepted original description (Kuhl 1820:103), the name is
given as Draco Timoriensis Pron. However, Pron never published a description of

a Draco from Timor, even though the specimens from his expedition were presented
to the Museum National dHistoire Naturelle in Paris. The name Draco Timoriensis
also appears in Dumril and Bibron (1837:454), who list a manuscript by Pron
first in their list of synonyms, with Kuhl (1820) relegated to second place. The
first mention of the name timorensis is probably an unjustified emendation by
Gray (1845), who listed the species as Draco Timorensis and referred to Draco vir-
idis Timorensis, a plate in Schlegel (183744). Subsequent authors, beginning with
Gnther (1864) and Boulenger (1885), have perpetuated this error even though
the latter corrected Gray in the spelling of the name attributed to Schlegel by listing
it as Draco viridis var. timoriensis. Since this change in the spelling of the specific
epithet is not via an accepted nomen substitutum (as suggested by Wermuth 1967),
the correct spelling for the flying lizard found on Timor remains Draco timoriensis.
Family Gekkonidae True Geckos

Genus Cyrtodactylus
http://species-id.net/wiki/Cyrtodactylus
Fig. 9
Common name. (E) Timor Bent-toed Gecko. *(T) Teki ain-fuan kleuk (teki = small
gecko, kleuk = bent, ain-fuan = toe).
Identification. This candidate species of Cyrtodactylus, designated as Cyrtodactylus
sp. 1 [Ca CMD 383], lacks the characteristic orange banding pattern of the tokay
gecko (Gekko gecko) and has dorsal patterning with a greater amount of brown compo-
nents (spots, flecks, lines) than any other gecko found on Timor. In its size, it is inter-
mediate between the common house geckos (Hemidactylus frenatus, Gehyra mutilata)
and the tokay gecko, and it does not have a flattened tail or dorsolateral skin flaps as in
H. platyurus. It is also the only gecko to possess non-dilated digits, unlike those found
in typical geckos. Instead the toes are slender and curved (Fig. 9), resulting in various
names being inconsistently applied to members of the genus (e.g., bent-toed geckos,
naked-toed geckos, bow-fingered geckos). The genus Cyrtodactylus is the most diverse
genus within the seven families comprising the Gekkota, with at least 130 species
described.
Collection and natural history. Two specimens of what is clearly an undescribed
species of Cyrtodactylus were captured on the same night at the Trilolo River near Same,
Manufahi District (altitude 553 m). There are substantial differences in pholidosis
and overall morphology from all known species of Cyrtodactylus (see Rsler and Glaw
2008: Table 1). One individual was found on a boulder-face along the riverbank, while
the second was in leaf litter at the foot of a large boulder at the boundary between
riverine habitat and coffee plantation.
Figure 9. Undescribed species of Cyrtodactylus. Female from near Same, Manufahi District (USNM
[CMD 383], SVL 58 mm, TL 127 mm). Photo by Hinrich Kaiser.
Gehyra cf. mutilata (Wiegmann, 1834)

http://species-id.net/wiki/Gehyra_mutilata
Fig. 10
Common names. (E) Mutilated Gecko, Stump-toed Gecko, Tender-skinned Gecko.

*
(T) Teki kulit kanek (teki = small gecko, kanek = injured, kulit = skin).
Identification. Individuals of the genus Gehyra (Fig. 10) in Timor-Leste are most
commonly seen around human habitations, where they occur sympatrically with the
common house gecko Hemidactylus frenatus. Identification on sight is usually quite diffi-
cult because of the superficial similarity of these two species. Upon capture, an early indi-
cation that a specimen is G. cf. mutilata is its ability to shed skin and scales as a defensive
mechanism. Unless great care is taken, the skin tears very easily at capture and the animal
will appear mutilated. Furthermore, the anterior and posterior postmental chin shields
are elongate and in broad contact down the midline in Gehyra mutilata, whereas in H.
frenatus these chin shields are shorter, more rounded, and only the anterior pair is in mid-
line contact, the posterior pair being widely separated by heterogeneous granular scales.
Collection and natural history. The three specimens of G. cf. mutilata we collected
occurred syntopically with Hemidactylus frenatus and were invariably collected at the same
time as specimens of that species. They occurred on the walls of houses as well as on the
trunks of trees. It is possible that G. cf. mutilata was introduced to Timor at some point
during prehistoric human colonization or pre-colonial or colonial inter-island trade.
Taxonomic comment. Even though G. mutilata sensu stricto is a widely distributed
species and occurs throughout Southeast Asia and the western Pacific realm, there is
very little known about its exact distribution in Wallacea (Fisher 1997). There are
several different names in the literature that could be applied to Gehyra populations
on Timor that are not mutilata. Until we unequivocally confirm the identity of our
specimens, they are here listed as G. cf. mutilata.
Figure 10. Gehyra cf. mutilata. Male (USNM [CMD 459], SVL 50 mm, TL 102 mm) from Lor 1 vil-
lage, Lautm District. Photo by Mark OShea.
Gekko gecko (Linnaeus, 1758)

http://species-id.net/wiki/Gekko_gecko
Fig. 11
Common names. (E) Tokay Gecko. (T) Toke.

The Tetun common names for geckos, teki (smaller geckos) and toke (the large to-
kay) are also used as slang meaning to identify young single women or men, respectively.
Identification. Tokay geckos are easily identified by their striking orange dorsal
patterning (Fig. 11), as well as by their aggressive open-mouth display when encoun-
tering a threat. They also have a distinctive vocalization (to-keh) that gave them their
common name. The dark-light banding pattern on the tail of hatchling G. gecko may
at first glance be confused with a similar pattern on the tails of some bent-toed geckos
(genus Cyrtodactylus). Based on overall habitat needs, if a gecko with a banded tail is
encountered on the walls of human habitations in Timor-Leste, it is most likely G.
gecko. Geckos of the genus Cyrtodactylus lack the dilated scansors necessary for climb-
ing walls and are generally not associated with man-made structures.
Collection and natural history. We found tokay geckos inhabiting nearly all of the
hotels and guest houses in which we stayed, in addition to many other buildings and
structures, as well as wooded regions in Lautm District (e.g., Lor 1 village). Tokay geckos
were not present at the higher altitude localities we searched (above 1000 m). We col-
lected one adult (Fig. 11 upper) and three juveniles (e.g., Fig. 11 lower) to secure vouchers
and then discontinued the collection of this species. Individuals were observed preying on
insects attracted by artificial light sources as well as on smaller geckos (e.g., Hemidactylus
Figure 11. Tokay gecko (Gekko gecko). Adult male from Same, Manufahi District (USNM 573671, SVL
142 mm, TL 236 mm; top) and juvenile G. gecko from Wailakurini, Viqueque District (USNM 573673,
SVL 88 mm, TL 168 mm; bottom). Note the brownish, regenerated tail on the adult (top). Photos by
Mark OShea (top) and Hinrich Kaiser (bottom).
frenatus). One particularly aggressive individual even attacked a smaller gecko that we
had stunned using a blowgun and placed into a plastic bag for safekeeping, and pulled it
behind a bamboo wall inside one of our sleeping cabins. Eggs of what we presumed to be
G. gecko from their size, and by the presence of adult tokays in the immediate vicinity of
the clutches, were discovered in communal groups in rotting logs, on the walls of huts,
and in tree holes. Based on their pattern of distribution and habits, it is possible that tokay
geckos were introduced to Timor via inter-island trading or during colonization.
Hemidactylus frenatus Schlegel, 1836

http://species-id.net/wiki/Hemidactylus_frenatus
Fig. 12
Common names. (E) Common Indo-Pacific House Gecko. *(T) Teki uma baibain
(teki = small gecko, uma = house, baibain = common).
Identification. See comments under Gehyra cf. mutilata.
Figure 12. Individuals of Hemidactylus frenatus showing two distinctive color patterns. (Top) Specimen
from near Baucau, Baucau District (USNM [CMD 526], SVL 47 mm, TL 90 mm) displaying a pattern
of distinct dorsolateral stripes complemented by an interrupted, less distinct vertebral stripe. Note the
regenerated tail and the bright orange mite infestation on the third toe, as well as an egg visible through
the skin. (Bottom) Specimen from near Lor 1 village (USNM [CMD 488], SVL 42 mm, TL 89 mm)
with a cryptic dorsal pattern. Photos by Mark OShea.
Collection and natural history. This species is the most commonly encountered
gecko in Asia, and it has also become an established exotic in many places in the New
World. As a perianthropic species, it is present on the walls or among the rafters of
almost every building, and it is distributed at widely differing altitudes and on the
edges of many different habitats. This species is among the several gecko species found
in Timor-Leste that may have been introduced during prehistoric colonization or his-
torical inter-island trade. The species appeared absent from pristine habitats such as
undeveloped forests, whereas it does occur in coffee plantations.
Hemidactylus platyurus (Schneider, 1792)

http://species-id.net/wiki/Hemidactylus_platyurus
Fig. 13
Common names. (E) Common Flat-tailed Gecko. *(T) Teki ikun belar (teki = small
gecko, belar = flat, ikun = tail).
Identification. Flat-tailed geckos can be identified by the presence of lateral skin
flaps and a flattened tail bearing a fringe of denticulate skin (Fig. 13). Such adaptations
provide an increased measure of cryptic morphology for these geckos in addition to
their bark-like coloration, as the skin extends almost seamlessly from body to substrate
and all but eliminates any shadow these animals may cast.
Collection and natural history. We collected several specimens by night when
they were exposed on the trunks of trees or under bark in forested habitats at elevations
Figure 13. Hemidactylus platyurus. Male from Lor 1 village, Lautm District (USNM [CMD 458], SVL
41 mm, TL 91 mm). Photo by Mark OShea.
of 300 m or less. In one instance, a pair was found in close proximity on the same tree.
Geckos like these may also be found in a perianthropic setting or in lowland savannas,
where they may be sympatric with H. frenatus and Gehyra cf. mutilata. As these other
two species, it is likely that H. platyurus is not native to Timor but was introduced by
human activities in the past.
Taxonomic comment. Until the revision of geckos in the genus Hemidactylus by
Carranza and Arnold (2006), this species was called Cosymbotus platyurus.
Family Scincidae Skinks

Genus Carlia
http://species-id.net/wiki/Carlia
Fig. 14
Common names. (E) Four-fingered Skinks. *(T) Mamr liman-fuan haat (mamr =
skink, haat = four, liman fuan = finger).
Figure 14. Representative specimens of the four presumed species of Carlia found in Timor-Leste. A
Carlia sp. 1, a high-altitude form from Maubisse, Ainaro District (USNM [CMD 361], SVL 37 mm, TL
99 mm) B Carlia sp. 3, a lowland form from near Lor, Lautm District (USNM [CMD 483], SVL 43
mm, TL 108 mm) C Carlia sp. 2, a highland form from the western versant of Mt. Ramelau in Ermera
District (USNM [CMD 401], SVL 44 mm, TL 112 mm) D Carlia sp. 4, a lowland form from near Bau-
cau, Baucau District (USNM [CMD 522], SVL 42 mm, TL 112 mm). Photos by Mark OShea.
Identification. Skinks of the genus Carlia are small lizars that are often found
foraging in grassy vegetation or under decaying palm fronds. Their identifying char-
acteristic is a four-fingered forefoot all other lizards in Timor-Leste possess five-
fingered forefeet. In common with all other lizards, they possess pentadactyl hindfeet.
Identification within the genus Carlia is often difficult and involves scale counts, color
patterns, morphometrics, and natural history characters. George Zug (USNM) pre-
liminarily verified the initial division of our specimens into groups.
Collection and natural history. We encountered four phenotypically distinct
four-fingered skinks of the genus Carlia (Fig. 14) throughout our survey. Considering
how morphologically conservative species in this genus are, we are as yet unable to
assign them to existing taxa with confidence. Comparisons with specimens of C. pero-
nii, C. spinauris, and C. fusca (including the holotypes of C. peronii and C. fusca and
a syntype of C. spinauris) show that the groups we found in Timor-Leste are similar
to the former based on overall body morphology and size. There are, however, differ-
ences with C. peronii and C. spinauris (fide Zug 2010) that require further study. We
acknowledge that a collection of Carlia in a relatively short time span may not allow us
to assess the true breadth of morphological variation since coloration may depend on
the degree of sexual maturity or reproductive readiness (G. Zug, pers. comm.). A more
detailed morphological and genetic analysis is therefore underway.
Among our Carlia specimens, we can differentiate two high-altitude forms and two
low-altitude forms. One of the high-altitude forms from the Maubisse and Same areas
(Ainaro and Manufahi District, respectively) on the eastern slopes of Mt. Ramelau (alti-
tudes 6001500 m) resembles species in the C. peronii group (sensu Zug 2010), and we
believe that this form may have previously been listed in the literature as C. peronii (e.g.,
Greer 1976). Based on our fresh material and on comparisons with preserved material,
we are more comfortable with listing this form as Carlia sp. 1 [Ca CMD 354] (Fig. 14A)
until we have completed a survey of material in herpetological collections. Similarly, a
second high-altitude form from the western versant of Mt. Ramelau in Ermera District
(altitudes near 1200 m) is also similar to species in the C. peronii group, and the orange
ventral coloration of sexually mature individuals makes it readily distinguishable (Fig.
14C). At this time, we prefer to list this form as Carlia sp. 2 [Ca CMD 400] until addi-
tional work is completed. This form was found exclusively during the daytime and near
dusk, in clearings at the edge of coffee plantations, amongst scattered piles of bamboo
husks, and on grassy areas surrounding bamboo stands. Several individuals were also
found in a pile of construction debris (metal piping and wooden boards). We noted the
absence of Carlia specimens in open spaces with no nearby form of cover (as measured
in a few lizard body lengths).
The two lowland forms are not easily placed in either of the above groups. Even
though their morphology is conservative, there are differences in overall gestalt and
coloration. Since our collections were conducted during what is considered a mainly
dry time of the year, we may see some changes when we return during the wet season,
as individuals may change color as they mature or reach reproductive readiness. At
this point, we consider the form from the coastal dry forest near Lor (Lautm Dis-
trict) as Carlia sp. 3 [Ca CMD 471] (Fig. 14B). The collection of a single specimen
from the Baucau area provides insufficient material to determine with clarity what
its specific status should be, but given its distinct morphology and pending the col-
lection of additional material we consider this form as Carlia sp. 4 [Ca CMD 522]
(Fig. 14D). At the Lor site, individuals were most predictably found by turning
over decaying palm fronds and in the leaf litter. In a typical display of skink behav-
ior, Carlia were seen each morning, basking, hunting, or displaying in the various
sunspots near our campsite.
Cryptoblepharus leschenault (Cocteau, 1832)

http://species-id.net/wiki/Cryptoblepharus_leschenault
Fig. 15
Common name. (E) Leschenaults snake-eyed skink. *(T) Mamr matan samea
(mamr = skink, matan = eye, samea = snake).
Identification. The defining characteristic of this genus, which occurs widely
across the tropical world, is the lack of moveable eyelids, which are replaced by
transparent snake-like brilles. Their slender body, long tail, and distinctive dorsal
pattern, consisting of two light dorsolateral stripes and a characteristic light mid-
dorsal line that forks posterior to the neck, easily identifies individuals of this species.
Our identification of this species was confirmed by reviewing the figures in Horner
(2007: Figs. 170, 171).
Collection and natural history. We collected four individuals of C. leschenault
and observed many others. These skinks were invariably seen on the trunks of hard-
woods above root level. Two species of Cryptoblepharus have been reported from
Timor, but we did not encounter C. schlegelianus, which can be distinguished from
C. leschenault by a greatly reduced degree of dorsolateral striping, particularly the
lack of a vertebral stripe that extends along the dorsum and onto the tail (Horner
2007: Fig. 188).
Taxonomic comment. Salomon Mller first mentioned the occurrence of this
species on Timor in letters written in 1829 (Brongersma 1942). Mller reported on
collections he made on Semau, a small island off the northwest coast of West Timor,
as well as on Timor. Mllers descriptions include C. schlegelianus and C. leschenault
and it appears that both of these species were collected within a short time span during
his visits. Based on the specimens available to Brongersma in the 1940s in the col-
lections at the RMNH (see Brongersma 1942) and our own search of herpetological
collections, there exists a single voucher specimen of C. leschenault from Timor in the
collection of the Naturhistorisches Museum Basel (NHMB 12885, as Ablepharus bou-
tonii leschenault), which was first reported by Forcart (1953). Our specimens therefore
comprise the first series of C. leschenault from Timor and Timor-Leste. Specimens col-
lected of this species by Max Weber (see Weber 1890) were from Flores (described as
Ablepharus boutonii furcata).
Figure 15. Cryptoblepharus leschenault. Male (USNM 573654, SVL 37 mm, TL 84 mm) from near Lor,
Lautm District. Photo by Mark OShea.
Figure 16. Individuals of two presumed undescribed species of Eremiascincus. A Eremiascincus sp. 1
(USNM [CMD 365], SVL 66 mm, TL 185 mm) from Maubisse, Ainaro District B Eremiascincus sp. 2
(USNM [CMD 474], SVL 51 mm, TL 101 mm) from Lor 1 village, Lautm District. Photos by Mark
OShea.
Genus Eremiascincus
http://species-id.net/wiki/Eremiascincus
Fig. 16
Common names. (E) Night Skinks. *(T) Mamr kalan (mamr = skink, kalan =
night).
Identification. Lizards of the genus Eremiascincus are relatively slender, long-bod-
ied skinks with rounded, elongate, conical tails and reduced limbs. On Timor it is
necessary to distinguish between at least five species of Eremiascincus (E. timorensis, E.
antoniorum, E. emigrans, Eremiascincus sp. 1 [Ca CMD 365], Eremiascincus sp. 2 [Ca
CMD 474]), and the distinctions between these are rather finite.
Collection and natural history. We collected two of the species of Eremiascincus
found on Timor. Eremiascincus sp. 1 (Fig. 16A) is a species whose distribution is appar-
ently limited to elevations above 1000 m. Individuals were primarily found under logs
and rocks, and were never encountered in the open during the day. Eremiascincus sp. 1
was the highest-altitude reptile we recorded in Timor-Leste (southwestern slopes of Mt.
Ramelau, 2046 m). We found three individuals of Eremiascincus sp. 2 (Fig. 16B) among
fallen and decaying palm fronds near Lor 1 village, Lautm District. These individuals
were found syntopically with many individuals of Carlia sp. 3. Their activity level was
highest just before dusk.
Taxonomic comment. In their recent comprehensive molecular study of desert
skinks (Australian members of Eremiascincus Greer, 1979), Mecke et al. (2009) re-
vealed that several species of the polyphyletic genus Glaphyromorphus Wells and Wel-
lington 1983, including the isolepis group sensu Greer (1990), required reassignment
to Eremiascincus. This expands the definition of Eremiascincus, previously a genus com-
prised entirely of desert forms, to one including tropical species. For these species,
which includes all named species from Timor as well as the candidate species listed
herein, we use the term night skinks since they appear to adopt a more nocturnal or
crepuscular activity cycle, in contrast to most other skink species.
Figure 17. Eutropis cf. multifasciata. Male (not captured, TL ca. 225 mm) from Lospalos, Lautm Dis-
trict. Photo by Stephen Richards.
Eutropis cf. multifasciata (Kuhl, 1820)

http://species-id.net/wiki/Eutropis_multifasciata
Fig. 17
Common names. (E) Common Sun Skink, Many-lined Sun Skink. *(T) Mamr loro
(mamr = skink, loro = sun).
Identification. Skinks of the genus Eutropis are among the most robust lizards in
Timor-Leste, following in size only Varanus timorensis and Gekko gecko. They can be
identified by their brown dorsal coloration, smooth scales, and brownish black lateral
blotches (Fig. 17).
Collection and natural history. We collected two specimens of a very robust
form of Eutropis that closely resembles E. multifasciata. Because of the limited
size of our sample and because of observable differences between specimens from
Timor-Leste and those from other parts of Southeast Asia, we report these speci-
mens as E. cf. multifasciata pending a more thorough morphological and genetic
analysis. Both specimens were collected during the day while foraging in grassy
vegetation.
Taxonomic comment. Until the genus Eutropis was proposed for Asian members
of the former circumtropical scincid genus Mabuya (Mausfeld and Schmitz 2003), this
taxon was known as Mabuya multifasciata.
Figure 18. Lamprolepis cf. smaragdina. Male (USNM 573669, SVL 92 mm, TL 216 mm) from Lor 1
village, Lautm District. Photo by Mark OShea.
Lamprolepis cf. smaragdina (Lesson, 1826)

http://species-id.net/wiki/Lamprolepis_smaragdina
Fig. 18
Common names. (E) Emerald Tree Skink. *(T) Mamr modok (mamr = skink, mo-
dok = green).
Identification. In individuals of L. cf. smaragdina that possess the name-giving
color pattern, identification is easy. There are no other lizards reported from Timor
whose anterior body coloration is an emerald green. Individuals that lack this color
pattern may be confused with Eutropis cf. multifasciata, although in a direct compari-
son the darker dorsal coloration and lateral spotting, along with stouter body propor-
tions, of the latter should be diagnostic. Individuals of Lamprolepis are generally found
on the upper portion of the trunk of trees and palms, just below the foliage or crown,
into which they will quickly retreat when disturbed during basking, whereas individu-
als of Eutropis were not observed on trunks at all and will retreat into grassy areas or
under ground-level cover.
Collection and natural history. The several specimens of L. cf. smaragdina we col-
lected were taken from the trunks of trees by blow-piping or hand-collecting. Several
individuals were observed basking in sunspots very close to a specific tree, to which they
retreated when disturbed. A retreat would usually occur in stages, first by climbing the
trunk of the apparent home tree to just below the trees foliage while the skink main-
tained visual contact with the intruding human and then, when the threat persisted, a
total retreat into the dense foliage or crown of the tree or palm. We were able to find
what we believe to be the same individuals of L. cf. smaragdina on the same tree during
several days of observation, indicating that these lizards display strong site fidelity.
Unlike the entirely emerald green L. smaragdina we have encountered elsewhere
(HK in Peninsular Malaysia, MOS and SJR in Papua New Guinea), the solid bright
green coloration of Timor-Leste specimens was limited to the anterior half of the body,
posterior to which it morphed into a beige brown with dark dorsal spots and dark lat-
eral striping (Fig. 18). Two of our specimens lacked any green coloration and sported
a beige brown, pepper-and-salt patterned dorsal coloration. Coloration was, surpris-
ingly, not sexually dichromatic, and among the two males and two females in our col-
lection both sexes are represented by one bicolor green and brown specimen and one
entirely brown specimen.
Taxonomic comments. The pepper-and-salt color pattern we observed in our
specimens is reminiscent of the patterns described for L. s. moluccarum by Barbour
(1911, 1912) and for L. s. elberti by Sternfeld (1920). We defer the decision on the
exact taxonomic allocation of our specimens until a more detailed comparison, to in-
clude the named Wallacean subspecies of L. smaragdina, has been conducted.
Genus Sphenomorphus
http://species-id.net/wiki/Sphenomorphus
Fig. 19
Common names. (E) Wedge skinks. *(T) Mamr ai laran (mamr = skink, ai laran =
forest).
Identification. The genus Sphenomorphus includes wedge skinks of greatly vary-
ing sizes and diverse morphologies. The superficially conservative morphology of these
skinks is contradicted by a significant number of differences in the details of scalation
and coloration. Beyond a recognition based on color pattern (as is straightforward for
the species shown in Fig. 19D), these forms are difficult to tell apart. Sphenomorphus
sp. 1 [Ca CMD 445] (Fig. 19A) possesses a series of paired dark paravertebral spots
running as two lines onto the tail. In both sexes, the ventral coloration is cream, and
males possess a black throat. Sphenomorphus sp. 2 [Ca CMD 356] (Fig. 19B) has a
more diverse pattern of spots on its back, including brown, golden, and black spots in a
complex arrangement. The throat is not black in males, and in both sexes the venter is
yellow. Sphenomorphus sp. 3 [Ca CMD 415] (Fig. 19C) has a dorsal color pattern that
is more uniformly brown, with some transverse golden dorsolateral striping. The venter
of both sexes is a dirty cream color. Sphenomorphus sp. 4 [Ca CMD 416] (Fig. 19D) is
easily differentiated from the other forms by its smaller size and by a characteristic black
lateral stripe that extends from the eye along the side of the body all the way to the tip
of the tail. Its dorsal coloration is more reddish brown than that of the other forms.
Collection and natural history. We collected four forms of Sphenomorphus at three
very distinct localities. In the area around Maubisse at altitudes >600 m, we encoun-
Figure 19. Representative examples of the four putative undescribed species of wedge skinks, genus
Sphenomorphus. A Sphenomorphus sp. 1 (USNM [CMD 446], SVL 58 mm, TL 150 mm) from Lor,
Lautem District B Sphenomorphus sp. 2 (USNM [CMD 364], SVL 66 mm, TL 185 mm) from Maubisse,
Ainaro District C Sphenomorphus sp. 3 (USNM [CMD 416], SVL 69 mm, TL 152 mm) from the Mele-
otegi River near Eraulo, Ermera District D Sphenomorphus sp. 4 (USNM [CMD 415], SVL 42 mm, TL
92 mm) from the same locality as the animal in C Photos by Mark OShea.
tered a highland form (Sphenomorphus sp. 2; Fig. 19B) that frequently shared its hiding
places with night skinks (Eremiascincus). A worker at a road construction site gave to
us one specimen smaller than typical individuals of Sphenomorphus sp. 2 but of a very
similar morphology. That lizard was already injured from rough handling and expired
shortly after we received it. Based on the surrounding vegetation, the altitude (>600
m), and several morphological features, we refer this specimen to Sphenomorphus sp.
2 pending the collection of additional specimens and a more thorough analysis. There
is superficial resemblance of Sphenomorphus sp. 2 to S. variegatus (Peters 1867), but a
further evaluation of museum specimens is necessary to verify any species assignment.
At a second highland locality, in the area of Eraulo (Ermera District) on the west-
ern side of the Mount Ramelau massif, we found two distinct forms of Sphenomorphus.
One of these, from the Meleotegi River, is a form with very distinctive dorsal pat-
terning (Fig. 19D). It was discovered while turning over flat rocks at the edge of the
river. Based on its morphology we document this form as Sphenomorphus sp. 4. In the
adjacent forest and plantation habitats we collected two specimens of Sphenomorphus
sp. 3 (Fig. 19C).
The lowland form (Fig. 19A) from the dry coastal forest at Lor (Lautm District)
is quite common throughout the habitat. Individuals are most easily found during the
daytime on the buttresses and roots of trees or whilst foraging in the leaf litter. This
form has strong resemblance to S. florensis (Weber 1890), but since that species cur-
rently has three subspecies aside from the nominate form (Dunn 1927: S. f. nitidus, S.
f. barbouri, S. f. weberi) additional comparative work is necessary to determine its exact

species affinity. It is also possible that the taxonomy is complicated by the possible
synonymy of S. florensis and S. melanopogon (Dumril and Bibron 1839); one of the
syntypes of S. melanopogon reportedly was collected on Timor. At this time, we prefer
to call this form Sphenomorphus sp. 1 (Fig. 19A).
Taxonomic comments. In her seminal work on the reptiles of the Indo-Australian
region, Nelly de Rooij (1915) provided species accounts for Lygosoma florense and L.
variegatus, having examined specimens of only the latter from Timor. Based on our own
examination of various type specimens, the Lesser Sunda species of the genus Sphenom-
orphus require careful additional investigation in order to confirm their species status and
distribution. A revision of S. florensis is currently being conducted by Glenn Shea (in litt.).
Figure 20. Varanus timorensis. Two adult males from dry coastal forest in Lor, Lautem District. Photos
by Mark OShea (top) and Hinrich Kaiser (bottom).
Family Varanidae Monitor Lizards

Varanus timorensis Gray, 1831
http://species-id.net/wiki/Varanus_timorensis
Fig. 20
Common names. (E) Timor Tree Monitor, Spotted Tree Monitor. (T) Lafaek rai-
maran (lafaek = crocodile or large lizard, rai = dirt, maran = dry). (Mambae) Loti.
(Fataluku) Puilolon.
Identification. Timor tree monitors are the largest lizards reported from Timor-
Leste. Their identifying color pattern consists of circular yellow ocelli that cover the en-
tire dorsum (Fig. 20 Upper). These lizards have nares that are positioned posterior to the
snout by nearly a third of the distance from eye to the tip of the snout (Fig. 20 Lower),
whereas they are positioned in close proximity of the snout in other lizards on Timor.
Collection and natural history. We captured, photographed, and released four
individuals of this small monitor lizard in the Lor area, Lautm District, and two in
the Tutuala Beach area (Pantai Walu). One individual was seen repeatedly on a road-
side retaining wall in Tutuala just west of the turnoff for Tutuala Beach. Another in-
dividual was observed basking in a sunspot in coastal dry forest from where it escaped
into the hole of a dead branch. Three individuals were active in the undergrowth in
coastal dry forest. Two individuals were hiding by night under loose bark on a tree and
could initially only be recognized by their exposed tails. We have also seen road-killed
specimens of this species on roadways running through rice paddies.
While driving along the coast road of northern Timor-Leste, we caught glimpses
of monitor lizards of uncertain species affinity crossing the blacktop. On at least one
occasion we were able to ascertain by visual identification that the individual was V.
timorensis, but on other occasions we could not make a positive identification. Varanus
indicus, a widespread Indo-Pacific species of monitor lizard with similar body aspect
and lifestyle to V. timorensis, has been reported from Timor, although we have so far
been unable to find any corroboration, either via photography, voucher specimen,
or artistic representation, that V. indicus is present on Timor. Such reports may stem
from fleeting identifications by visitors familiar with the V. indicus body morphology.
Reports from local villagers regarding the presence of a large monitor lizard similar to
V. salvator from swamps in the Becora area require investigation.
Snakes (Order Serpentes)

Our current treatment of the snakes of Timor-Leste is primarily restricted to the land-
inhabiting species. Exceptions are those species, such as sea kraits (Laticauda) or file-
snakes (genus Acrochordus), whose lifestyle frequently places them on land or into land-
associated aquatic ecosystems. However, eleven species of seasnakes (Acalyptophis peronii*,
Aipysurus apraefrontalis*, A. duboisii*, A. foliosquama*, A. fuscus*, A. laevis*, Astrotia stokesii,

Emydocephalus annulatus*, Hydrophis melanocephalus, Lapemis hardwickei, Pelamis platu-
rus) have been collected from the reefs of the Sahul Shelf to the southwest of Timor
(Hibernia, Ashmore, Cartier and Scott Reefs; Minton and Heatwole 1975). One of us
(MOS) visited Hibernia and Ashmore Reefs in 2001 and recorded the seven species
marked by asterisks above, although A. apraefrontalis, A. foliosquama, and A. fuscus are be-
lieved endemic to the reefs. The other species may easily occur along the coasts of Timor,
on coral reefs, in estuaries or in coastal waters, as may species distributed further east on
the Australian and New Guinea coasts, or further north in the Banda or Flores Seas.
Family Colubridae Typical Snakes

Coelognathus subradiatus (Schlegel, 1837)
http://species-id.net/wiki/Coelognathus_subradiatus
Fig. 21
Common names. (E) Timor Racer, Lesser Sunda Racer, Lesser Sunda Trinket Snake.
*
(T) Samea laho (samea = snake, laho = rat).
Identification. Coelognathus subradiatus is a slender brown racer that can be iden-
tified by a pair of black paravertebral stripes that run along its back from the rear of
Figure 21. Coelognathus subradiatus. Male from Baucau town, Baucau District (USNM 573676). Photo
by Mark OShea.
the head to the tail, with varying degree of completeness (Fig. 21). This snake also has
a short (13 scales) longitudinal postorbital stripe and a flat, squared tip to the snout.
Collection and natural history. One individual was captured at night on the
road leading down from Old Town Baucau into the surrounding lowlands, but still in
the outskirts of the city (altitude 350 m). It had been hit by another vehicle and suc-
cumbed to its injuries shortly after collection. Coelognathus subradiatus populations in
Timor-Leste may not be conspecific with those on Roti, a neighboring island to the
west, or from some of the other Lesser Sunda Islands (Monk et al. 1997).
Taxonomic comment. We follow the taxonomy of Schulz (1996) who places the
populations of C. subradiatus from Timor, the type locality according to the description
of specimens by Schlegel (187344) collected there by Salomon Mller, in his Group
1. Of interest is that a subspecies of C. subradiatus was designated by Bethencourt Fer-
reira (1897) for a population from Timor, described as Coluber melanurus (Schl.), var.
timoriensis, n. var. from specimens in the Museo Bocage in Lisbon. Whereas Coluber
melanurus Schlegel 1837 is currently known as Coelognathus flavolineatus, Bethencourt
Ferreiras (1897, 1898) detailed description clearly allows identification of the Lisbon
specimens as C. subradiatus. We believe that in his description, and lacking a familiarity
with actual specimens of C. subradiatus from other collections, Bethencourt Ferreira
(1897, 1898) erred on the side of caution by identifying his specimens as a unique vari-
ant related to Coluber melanurus and young Coluber erythrurus (Dumril et al. 1854).
Bethencourt Ferreiras accounts were apparently cited only twice in relation to this snake
population, first by Barbour (1912) and then by de Rooij (1917). Without comment,
Barbour (1912:195) lists this population as a full species (Elaphe timoriensis) in his
expansive table, considered an error by Schulz (1996). In an unfortunate set of circum-
stances, this confused situation has been compounded by the complete loss of speci-
mens and collection data in the Museo Bocage by fire in 1978. Until the revision of Old
World ratsnakes by Helfenberger (2001), this species was placed in the genus Elaphe.
Dendrelaphis inornatus timorensis (Smith, 1927)

http://species-id.net/wiki/Dendrelaphis_inornatus_timorensis
Fig. 22
Common names. (E) Timor Bronzeback, Timor Treesnake. *(T) Samea kotuk kr kaf
(samea = snake, kotuk = back, kr kaf = brown).
Identification. Bronzebacks are slender, diurnal snakes capable of rapid arboreal
locomotion that may confuse the eye. The Timor bronzeback is brown above and with
a greenish cream venter (Fig. 22 Upper). A narrow black stripe separates the dorsum of
the head from the paler labial scales of the mouth (Fig. 22 Lower). When threatened,
bronzebacks may inflate their neck, exposing the blue interstitial skin between their
scales and making themselves look larger and more threatening to potential attackers.
Identification and comparison with the widespread common bronzeback Dendrelaphis
pictus was made in accordance with How et al. (1996) and David and Vogel (1996)
and was verified by Gernot Vogel. The taxonomy of D. inornatus is currently undergo-
ing a re-evaluation by Vogel and Jan van Rooijen (Gernot Vogel, pers. comm.).
Collection and natural history. We collected three specimens of this subspecies
(one adult, two juveniles). Two were found at night in a resting position in shrubs
or bushes no higher than 2 m off the ground. These sleeping snakes became alert
once illuminated by our flashlights, and they attempted to escape thereafter. The third
specimen was observed while it travelled through the leaf litter in dry coastal forest. A
fourth specimen was seen in the proximity of the third, but on the trunk of the tree.
When pursued, this snake rapidly ascended the trunk and disappeared in the foliage.
Figure 22. Dendrelaphis inornatus timorensis. Male (USNM [CMD 493], SVL 689 mm, TL 1054 mm)
from Lor 1 Village, Lautm District. Photos by Hinrich Kaiser.
Lycodon capucinus (Boie, 1827)

http://species-id.net/wiki/Lycodon_capucinus
Fig. 23
Common names. (E) Common Wolfsnake. (T) Samea lobo (samea = snake, lobo = wolf).
Identification. Common wolfsnakes have a dorsally brown body with a series of
weak pale yellow to white bands (Fig. 23). The dorsal part of the head is uniformly
brown, offset from the rest of the body by a pale yellow nuchal band. The labial scales
and venter are cream colored.
Collection and natural history. Our single specimen of L. capucinus was collected
by local people in the town of Same (Manufahi District) while crossing the towns main
road after a heavy rain. It was brought to us undamaged in a 500-ml clear plastic water
bottle.
Taxonomic comment. In the Lycodon literature the names L. aulicus and L.
capucinus are seemingly used interchangeably, sometimes with capucinus relegated to
subspecific status within aulicus. During the time when capucinus had subspecific sta-
tus, some authors did not differentiate it from aulicus sensu stricto. When capucinus
was controversially re-elevated to specific status, this compounded an already confus-
ing situation. As a consequence, the Lycodon forms from Southeast Asia and Wallacea
under consideration here have been known by three possible species and subspecies
Figure 23. Lycodon capucinus. Male (USNM 573681, SVL 395 mm, TL 491 mm) from the town of
Same, Manufahi Distict. Photo by Mark OShea.
names. We here follow Taylor (1965) and David and Vogel (1996) in using the name
L. capucinus for the Lesser Sunda form.
Lycodon subcinctus Reinwardt, 1827

http://species-id.net/wiki/Lycodon_subcinctus
Fig. 24
Common names. (E) Malayan Banded Wolfsnake. *(T) Samea kadeli (samea = snake,
kadeli = ring).
Identification. Banded wolfsnakes are easily identified by the series of contrast-
ing white to cream-colored bands that offset the dark brown to black body coloration
(Fig. 24). In this coloration, they mimic Malayan or many-banded kraits (Bungarus
candidus and B. multicinctus, respectively), highly venomous species, with which this
wolfsnake is sympatric in northern parts of its range.
Collection and natural history. In contrast to the careful capture of the Lycodon
in Same, our specimen of L. subcinctus was obtained within minutes of having been
hacked to death at an elementary school. The animal had reportedly been found in
the school and was disposed of just as we explained our purpose to some of the local
residents. The snake was handed to us draped dead over a branch, with body segments
merely attached by threads of skin.
Figure 24. Lycodon subcinctus. Damaged specimen (USNM 573682, SVL 544 mm, parts of tail lost)
from Letefoho, Manufahi District. Photo by Mark OShea.
Family Homalopsidae Oriental and Australasian Mudsnakes

Cerberus cf. rynchops (Schneider, 1799)
http://species-id.net/wiki/Cerberus_rynchops
Fig. 25
Common names. (E) Bockadam, Dog-faced Watersnake. *(T) Samea natar (samea =
snake, natar = rice paddy).
Identification. Dog-faced watersnakes are easily distinguished based on both mor-
phology and habits. Compared with other snakes, the head is blunt with a rounded
snout and relatively small eyes in a dorsolateral position (Fig. 25). Coloration is brown-
ish gray, often with darker blotches on the back. Relative to most snakes in Timor-Leste,
these snakes are heavy-bodied (body diameter is robust as opposed to slender). These
snakes are most frequently encountered in habitats with standing or slow-flowing water.
Collection and natural history. We captured a single specimen of a snake very sim-
ilar to C. rynchops at night from a flooded rice paddy near Baucau (Baucau District). The
snake submerged when illuminated, but was extracted from the muddy water with ease.
Taxonomic comment. In its overall morphology our specimen clearly resembles
C. rynchops, but a few specific characteristics of its scalation are intermediate between
C. rynchops and C. australis. Until we are able to ascertain its precise taxonomic status,
through more detailed morphological and genetic comparisons, we list this specimen
Figure 25. Cerberus cf. rynchops. Male (USNM 573675, SVL 598 mm, TL 756 mm) from a rice paddy
in the Baucau area, Baucau District. Photo by Mark OShea.
as C. cf. rynchops. The genus Cerberus is currently undergoing a taxonomic evaluation

by John C. Murphy (in litt.).
Family Typhlopidae Blindsnakes

Ramphotyphlops braminus (Daudin, 1803)
http://species-id.net/wiki/Ramphotyphlops_braminus
Fig. 26
Common names. (E) Brahminy Blindsnake, Flowerpot Snake. *(T) Samea matan
delek (samea = snake, matan delek = blind).
Identification. Brahminy blindsnakes are vermiform snakes in both morphology
(Fig. 26 Upper) and behavior. Body thickness is similar in diameter to the ink tube of
a ballpoint pen, and when encountered these snakes will writhe energetically. When
grabbed, a typical behavior is to stab the pointed end of the tail into the finger hold-
ing the animal in order to gain better purchase for an escape. A closer look will reveal
much-reduced eyes as pigmented areas under translucent head scales (Fig. 26 Lower),
a tiny forked tongue, and a scale pattern that is diagnostic when differentiating blind-
snake taxa. Addison Wynn (USNM) confirmed species identity.
Collection and natural history. We found three specimens of this near-cosmo-
politan blindsnake, each in disturbed habitat. The first was found under a rock in the
middle of an unpaved country lane with very little vehicular traffic. The second was
spotted within minutes of the first under a rock along the edge of the same road. We
were surprised by fact that the third specimen essentially found us, by travelling across
the smooth, tiled floors of the hotel lobby and into one of our rooms. Even though it
was easily spotted, it was quite difficult to pick up.
This is the only known parthenogenetic snake species and this factor, combined
with its small size and secretive nature, make it an excellent colonizer. A single speci-
men arriving in the root-ball of a decorative or food plant is sufficient to establish a
new colony. Due to the actions of humans this is the most widely distributed snake
species in the world, probably only rivaled by the ubiquitous house geckos (Hemidac-
tylus spp.) amongst the lizards.
Family Viperidae True Vipers and Pitvipers

Cryptelytrops insularis (Kramer, 1977)
http://species-id.net/wiki/Cryptelytrops_insularis
Fig. 27
Common names. (E) Lesser Sunda Island Pitviper, Island Pitviper, Lesser Sunda
White-lipped Pitviper. (T) Samodok. (Mambae) Samor. (Fataluku) Cuale.
Figure 26. Ramphotyphlops braminus. Specimen (USNM 573683, SVL 147 mm, TL 150 mm) from the
Same area, Manufahi District. Photos by Hinrich Kaiser (top) and Mark OShea (bottom).
Figure 27. Cryptelytrops insularis. Female (USNM [CMD 594], SVL 684 mm, TL 784 mm) from the
flood plain of Lake Ira Lalaro, Lautm District. Photo by Hinrich Kaiser.
Identification. This snake can be distinguished by both morphology and typical

behavior. Characteristic of a pitviper are the paired forward-facing, heat-sensitive pits,
posterior to and below to the smaller lateral-facing nares, and the vertically elliptical
pupils of the eyes (Fig. 27). Pitvipers documented from Timor-Leste are most fre-
quently bright green in dorsal color with the exception of a reddish stripe on the poste-
rior-most portion of the tail. There exists a second, yellow color morph that apparently
is seen in low-rainfall areas in Timor-Leste (CRT, pers. obs.) and also on some of the
neighboring islands. Specimens from Wetar are bright yellow whilst some of those
from the Komodo Islands are cyan. In the green morph, the labial scales are a yellowish
green. The characteristic behavior of these snakes when threatened is to coil the body
tightly (Fig. 23) for a defensive strike. These are currently the only venomous reptiles
confirmed from Timor-Leste whose bite may have serious implications for humans.
Collection and natural history. We obtained four specimens of this arboreal pitviper
in four localities with distinct habitat types, all in eastern Timor-Leste (Baucau, Viqueque,
and Lautm Districts). It is remarkable to note that none of the individuals we located
were active in trees, shrubs, or leafy vegetation, as might be expected of members of a
supposedly arboreal genus, but were located exclusively on the ground. This leads us to
speculate that this island form of the widespread green pitviper ecomorph could be less
arboreal than some of its congeners and more of a habitat generalist in the absence of com-
petition from terrestrial pitvipers or true vipers, such as the Southeast Asian Russells viper
(Daboia siamensis) that occurs in sympatry with this species further northwest in the Lesser
Sundas. The first specimen we collected was a recent road-kill on the Baucau-Lautm road
and was found during the daytime within a very short distance of the beach in dry coastal
forest. All other specimens were found by night. Our attention was drawn to the second
specimen near Timor Village Hotel by a hotel-worker, who encountered it while walking
home in an area of short grass adjacent to human habitations. The third specimen (Fig.
23) was encountered while it rested on a bed of decaying foliage in a forested flood plain
adjacent to the large open Lake Ira Lalaro flood plain. The last specimen was found at the
edge of a dry rice paddy on the outskirts of Baucau. There were no trees within ca. 50 m of
this snakes position. Reports indicate that pitvipers are relatively common in grassy areas
and agricultural plots near human habitations, creating a dangerous situation for barefoot
humans active during the hours of darkness. The potential snakebite risk posed by this
species is being investigated in collaboration with medical colleagues.
Taxonomic comment. This species was regarded as a subspecies of the widespread
Asian species Trimeresurus albolabris until it was elevated to full species rank by Gian-
nasi et al. (2001). Malhotra and Thorpe (2004) changed the generic assignment from
Trimeresurus to Cryptelytrops in their revision of Asian pitvipers.
Turtles (Order Testudines)

Our treatment of the turtles of Timor-Leste is primarily concerned with terrestrial spe-
cies and those living in land-associated aquatic ecosystems. However, six of the worlds
seven sea turtle species occur in the northern coastal waters of Australia (Caretta caret-
ta, Chelonia mydas, Dermochelys coriacea, Eretmochelys imbricata, Lepidochelys olivacea,
Natator depressus) and these will be recorded as the opportunity arises. We serendipi-
tously observed and filmed a sea turtle, most likely an Olive Ridley (L. olivacea), in the
surf of the Timor Sea off the beach at Lor 1 village (Lautm District). Bleeker (1860)
reported specimens of C. mydas and E. imbricata from Wetar Strait.
Family Chelidae South American and Australasian Side-necked Turtles

Chelodina timorensis McCord et al., 2007
http://species-id.net/wiki/Chelodina_timorensis
Fig. 28
Common names. (E) Timor Snake-necked Turtle. *(T) Lenuk kakorok ular (lenuk =
turtle, kakorok = neck, ular = snake).
Identification. Snake-necked turtles are easily distinguished by their long serpen-
tine necks (Fig. 28 Lower), which are fully as long as the entire carapace and which
permit the turtle to reach anywhere on its body.
Figure 28. Chelodina mccordi timorensis. This specimen was presented to us by a resident of Malahara
village, Lautm District. The lower panel shows how this turtle can bend its neck under its carapace when
threatened. Photos by Hinrich Kaiser.
Collection and natural history. During our survey we had heard that local villagers
occasionally keep live specimens of this protected snake-necked turtle as status symbols
or for trade, a clear contravention of the CITES protocols and an important reason
why the government of Timor-Leste is considering acceding to the CITES treaty. The
turtle has a highly restricted distribution in Lake Ira Lalaro near the easternmost point
of Timor-Leste. The lake itself is primarily seasonal, with water exiting the lake through
the Irasequiro River. The river itself does not reach the ocean but disappears beneath an
extensive limestone karst escarpment, the Paitxau Range (max. elevation at Mt. Paitxau,
925 m). During our visit we inquired about the availability of a snake-necked turtle for
photography, and we learned about a turtle that could be photographed. We declined
to purchase the turtle but offered US$ 5 for being given the opportunity to take pho-
tographs, in a symbolic gesture and specifically to prove that a living turtle could real-
ize revenue without being traded. This population is closely related to the Roti Island
Snake-necked Turtle (C. mccordi; Kuchling et al. 2007; McCord et al. 2007).
Taxonomic comment. Even though McCord et al. (2007) described the Lake
Ira Lalaro snake-necked turtle population as a distinct species, this decision was not
without controversy (see Kuchling et al. 2007), and the new species was not recog-
nized as an acceptable name by the CITES committee. However, the population is still
protected as an endangered species under CITES Appendix II. Kuchling et al. (2007)
described this population as C. mccordi timorlestensis.
Family Geoemydidae South American and Asian Pond Turtles

Mauremys reevesii (Gray, 1831)
http://species-id.net/wiki/Mauremys_reevesii
Fig. 29
Common names. (E) Reeves Stripe-necked Turtle, Chinese Pond Turtle. *(T) Lenuk
kakorok riskadu (lenuk = turtle, riskadu = striped, kakorok = neck).
Identification. Chinese pond turtles are readily identified by the characteristic yel-
low striping and blotching on their necks (Fig. 29).
Collection and natural history. The staff at the Albergaria Planalto, New Town
Baucau, became aware of our purpose in collecting specimens of amphibians and rep-
tiles and showed us an unidentified turtle that was kept in a small, stone-encased pond
on the grounds of the property. We were told that there were three turtles like this in
the area, one collected near Dili, and the two others just across the street in an empty
lot at the edge of town. Two of these had escaped by the time of our arrival, but we
were able to obtain this specimen as a voucher. The presence of M. reevesii was briefly
mentioned by McCord et al. (2007), and we reported details regarding its presence in
Timor-Leste elsewhere (Kaiser et al. 2010).
Figure 29. Mauremys reevesii. Male individual of the introduced Chinese pond turtle from the Albergaria
Planalto in New Town Baucau, Baucau District. Photo by Mark OShea.
Crocodiles (Order Crocodilia)

Family Crocodylidae Crocodiles
Crocodylus porosus Schneider, 1801

http://species-id.net/wiki/Crocodylus_porosus
Fig. 30
Common names. (E) Saltwater Crocodile, Indo-Pacific Crocodile, Estuarine Croco-

dile, Naked-necked Crocodile. (T) Lafaek tasi (lafaek = large lizard, tasi = ocean).
Identification. As the only crocodilian known from Timor and as a creature at the
root of the Timorese creation myth, this species probably requires no detailed descrip-
tion. However, C. porosus differs from other crocodiles in the possession of less dermal
armor, including lacking the typically four crocodilian post-occipital scutes (between
the rear of the skull and a cluster of six nuchal scutes over the shoulders) so obvious on
the necks of other species, hence the name naked-necked crocodile.
Collection and natural history. We saw and photographed several individuals
of C. porosus in the wild, most frequently in rivers while driving across bridges. We
also saw two captive individuals in Aileu, Aileu District. There exists a substantial
population of C. porosus in Lake Ira Lalaro, comprising several hundred individuals
with sizes exceeding 3 m in total length (CRT, pers. obs.). Crocodiles are commonly
reported from the swamps and swamp forests along the south coast of Timor-Leste,
Figure 30. Crocodylus porosus. Captive specimen in an enclosure in Aileu, Aileu District. Photo by Hin-
rich Kaiser.
which is the area with the greatest frequency of reported crocodile attacks. Our lim-
ited observations on the status of crocodiles in Timor-Leste have been published else-
where (Kaiser et al. 2009).
Discussion
The emergence of Timor-Leste as an independent nation comes at a precarious time for
its natural environment. During the centuries of colonialism and during over two dec-
ades of Indonesian occupation, a neglect of environmental management has led to the
unsustainable exploitation of natural resources for short-term gain and to the concomi-
tant indiscriminate destruction of habitats. This is perhaps nowhere more strikingly
seen than in the disappearance of the famed white sandalwood (Santalum album) forests
throughout most of Timor-Leste (McWilliam 2005). Even though the new country is
including environmental policy as a priority of nation-building, the damage sustained
by the infrastructure in the aftermath of the 1999 independence referendum, the lack of
capacity to educate about environmental issues, and the near lack of economic activity
in many areas of the country place a heavy burden on an already depleted environment
(see Sandlund et al. 2001 for a review). Furthermore, the factors traditionally listed
when outlining pressures on the environment in tropical developing countries (natural

or human-caused habitat loss, soaring population growth, economic need; Bradshaw et
al. 2009 and references therein) all apply to newly pacified Timor-Leste.
Perhaps against any expectations of undocumented diversity, our relatively limited
survey already shows a level of biodiversity for Timor-Leste that is higher than reported
previously and includes considerable endemicity. The occurrence of a species of Cyr-
todactylus extends the range of this genus to its southeastern extreme. Skinks of the
genus Carlia appear in a morphological array that constitutes several new species, akin
to some Carlia hotspots in Australia and New Guinea (e.g., Couper et al. 2005) with
their substantial cryptic diversity. Wedge skinks (genus Sphenomorphus) are a diverse
group throughout Southeast Asia, but their high diversity on a single island is note-
worthy. Lastly, the diversity of night skinks (genus Eremiascincus) appears to have been
underestimated on Timor and elsewhere (e.g., Mecke et al. 2009).
Timor-Leste is not unique in this juxtaposition of extensive environmental degra-
dation and a substantial number of undescribed species. This is certainly a Southeast
Asian theme (e.g., Giam et al. 2010) but it can be observed in most tropical environ-
ments, but especially on some of the nearby islands of Wallacea. Just as in other coun-
tries, in which high biodiversity has been discovered despite severe habitat degradation
(e.g., islands of the Caribbean: Smith et al. 2005), Timor-Leste poses a conundrum:
while biodiversity is high, it is in peril. It is fortunate that the Government of Timor-
Leste has established agencies that promote environmental education, the establish-
ment of natural parks and protected zones, and the establishment of sustainable use
practices in rural development, and we are hopeful that the threat to the remaining
pristine habitats and to the countrys biodiversity can be addressed and managed.
With a view to the realities of human-environment interactions, it is perhaps note-
worthy that many of the new species we list appear to possess sufficient phenotypic
plasticity to survive or even thrive in degraded habitats. For example, our observations
of rice paddy frogs include towns, active and fallow rice paddies, and forested areas. At
least one form of night skinks, wedge skinks, and four-fingered skinks occur in, or in
close proximity to, villages and their subsistence agricultural plots. The only new species
apparently limited to a more pristine or secluded environment was the bent-toed gecko,
but even it was collected in a transition zone, between a riverbed and a coffee plantation.
In the habitats we surveyed (e.g., Fig. 2), we were sometimes able to observe a very
precise division of ecological niches. For example, in the dry coastal forest in Lor, Lautm
District, six different species of lizards (four skinks, the flying lizard, and the monitor
lizard) were seen in syntopy. The monitor lizard was seen foraging in the leaf litter, among
the groundcover, and also on tree trunks, and among these six species it appears to be
the most generalized in its foraging habits. Careful observation of the five other species
quickly showed that microhabitat partition existed. Whereas Carlia would primarily for-
age among the dead, dry leaves, in a much more restricted fashion than Varanus timorensis,
Sphenomorphus would occupy the bases, exposed roots, and buttresses of trees. Individuals
of Cryptoblepharus were usually seen at intermediate heights on the trunks of trees, from
14 m high. Some individuals were also seen on thinner branches leading away from the
main trunk at comparable heights above the ground. Specimens of Lamprolepis occupied
the highest position among skinks, usually very close to the transition between the bare
trunk and the trees foliage or crown. Above the skinks we observed displaying males of
Draco whose capacity to glide makes this highest position the most desirable.
The results of this survey have been startling as an academic pursuit but also as an
historical and cultural journey. The lingering effects of a valiant struggle for independ-
ence can temper the excitement of discovering new species, as some of the specimens
we collected may have shared their habitat with freedom fighters and their local sup-
porters less than a decade earlier. The interest of both the government of Timor-Leste
and the countrys culturally diverse population in supporting those who seek to learn
about these habitats has been singularly rewarding and is a positive sign for sustain-
ability and conservation efforts.
Acknowledgments
First and foremost, we wish to acknowledge the tremendous support we received from
His Excellency, Prime Minister Xanana Gusmo of Timor-Leste with whose explicit writ-
ten permission this research was carried out. Without his intervention, and without the
invaluable and consistent help of his advisor Claudia Abate, we would not have been
able to start this project. We would also like to thank Manuel Mendes (Director of Na-
tional Parks) and Augusto Pinto (Director of Environment) for their understanding and
patience, and for their support of our research. SJR is grateful to Leonisa Lobato, Vasco
Leitao, Jorge Rui de Carvalho Martins (Division of Environment) and Cesar Jose da Cruz
(Director General, Ministry of Agriculture, Forestry, and Fisheries) for their assistance
and approval of export permits. For their capable and enduring assistance in the field
we thank Margaret Andrews, Kris Helgen, Luis Lemos, Dan Polhemus, Agivedo Laca
Ribeiro, Zito Afranio, and Annie Suzio-Granett. We thank Francisco da Costa Araujo for
presenting us with the specimen of Lycodon capucinus and Paul da Silva for the specimen
of Mauremys reevesii. Without the tireless assistance of Pedro Mesquita with transporta-
tion, we might have been stranded in Same. Additional logistics support was provided
by Anders Hofstee (Bakhita Mission), Paulo Aniceto (Rentl), Paul da Silva (Albergario
Planalto), Jeff Collins (Same), and Austasia Airlines and Turkish Airlines. We particu-
larly thank Roy McDiarmid for facilitating our research by supplying key literature, Steve
Gotte for assisting with specimen deposition, and George Zug for sharing his expertise
on the genus Carlia. Aaron Bauer helped with the identification of geckos, Mark Auliya
assisted with the identification of Mauremys reevesii, and Gernot Vogel identified the Den-
drelaphis. Glenn Shea provided some insights into the diversity and taxonomy of Lesser
Sunda wedge skinks. Peter Uetz made some very useful comments on the nomenclature
of Draco timoriensis, Jimmy McGuire commented on the presence of Draco in the Lesser
Sunda Archipelago, and John Murphy shared his expertise and recent research on snakes
in the genus Cerberus. Jean Lescure (MNHN) provided some key information regard-
ing the history of explorations in the Lesser Sunda Archipelago. The assistance of Steve
Gotte (USNM), Darrel Frost (AMNH), David Kizirian (AMNH), Annemarie Ohler
(MNHN), Gunther Khler (SMF), Pim Arntzen (RMNH), Ronald de Ruiter (RMNH),
and Raffael Winkler (NHMB) in allowing HK to examine specimens under their care is
acknowledged with much gratitude. Fieldwork was supported by a Title V Grant to Victor
Valley College (VVC), by three VVC Associated Student Body grants to JC, SH, and CS,
and by a grant from the American Society of Mammalogists to K. Helgen. This is Com-
munication 4 of the Tropical Research Initiative at Victor Valley College.
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Appendix I. Specimens Examined

In the following enumeration, specimens are listed using the collection acronym and
a specimen number. The specimen number is composed either of numbers only, or of
capital letters followed by numbers. The letters indicate the field numbers of Stephen
J. Richards (SJR) or Christine M. Kaiser (CMD). Specimens with field numbers have
not been accessioned yet at their respective depositories, but the specimens will never-
theless be accessible via a database search of field numbers at these institutions.
Frogs and Toads

Family Bufonidae
Duttaphrynus melanostictus.SAMA [SJR 7127], USNM 53765053
Family Dicroglossidae
Fejervarya sp. 1 [Ca CMD 431].USNM [CMD 429, 431, 43536, 508, 510,
51617, 538, 54041, 54345].
Fejervarya sp. 2 [Ca CMD 507].USNM [CMD 410, 426, 43334, 50607,
51215, 519, 53336, 539, 542].
Fejervarya sp. 3 [Ca CMD 355] .USNM [CMD 355, 382, 392, 428, 430, 432,
46970, 509, 511, 518, 568].
Limnonectes timorensis.SAMA [SJR 7072, 707578, 709293, 7099100,
710810, 711213, 7117, 7122, 712830, 713439], USNM [CMD 422
23].
Family Hylidae
Litoria everetti.USNM [CMD 42021].
Family Rhacophoridae
Polypedates cf. leucomystax.SAMA [SJR 707374, 707982, 710206, 7118
21, 713133, 7140], USNM [CMD 35759, 381, 419, 427, 457, 46467,
497502].
Lizards
Family Agamidae
Draco timoriensis.USNM 57365863.
Family Gekkonidae
Cyrtodactylus sp. 1 [Ca CMD 383].USNM [CMD 38384].
Gehyra cf. mutilata.SAMA [SJR 707071], USNM [CMD 45960, 505].
Gekko gecko.SAMA [SJR 709495], USNM 57367174.
Hemidactylus frenatus.USNM [CMD 441, 47678, 48890, 52426].
Hemidactylus platyurus.USNM [CMD 43738, 458, 494, 50304].
Family Scincidae
Carlia sp. 1 [Ca CMD 354].USNM [CMD 354, 36063, 36869, 371, 403,
406, 41213].
Carlia sp. 2 [Ca CMD 400].USNM [CMD 40002, 40405, 40708, 411,
414].
Carlia sp. 3 [Ca CMD 471].USNM [CMD 47173, 48385].
Carlia sp. 4 [Ca CMD 522].USNM [CMD 522].
Carlia incertae sedis.SAMA [SJR 700104, 706869, 708586, 7098, 7116,
712325.
Cryptoblepharus leschenault.USNM 57365457.
Cryptoblepharus incertae sedis.SAMA [SJR 708384
Eremiascincus sp. 1 [Ca CMD 365].USNM [CMD 36567, 370, 37274,
38588, 39399].
Eremiascincus sp. 2 [Ca CMD 474].USNM [CMD 474, 48687].
Eremiascincus incertae sedis.SAMA [SJR 705667, 7097, 7107, 711415, 7126].
Eutropis cf. multifasciata.SAMA [SJR 708891, 7096], USNM 573664666.
Lamprolepis cf. smaragdina.USNM 57366770.
Sphenomorphus sp. 1 [Ca CMD 445].USNM [CMD 44549, 45155].
Sphenomorphus sp. 2 [Ca CMD 356].USNM [CMD 356, 364].
Sphenomorphus sp. 3 [Ca CMD 415].USNM [CMD 415].
Sphenomorphus sp. 4 [Ca CMD 416].USNM [CMD 416].
Sphenomorphus incertae sedis.SAMA [SJR 7111].
Snakes
Family Colubridae
Coelognathus subradiatus.USNM 573676.
Dendrelaphis inornatus timorensis.USNM [CMD 450, 468, 493].
Lycodon capucinus.SAMA [SJR 7087], USNM 573681.
Lycodon subcinctus.USNM 573682.
Family Homalopsidae
Cerberus rynchops.USNM 573675.
Family Typhlopidae
Ramphotyphlops braminus.USNM 57368385.
Family Viperidae
Cryptelytrops insularis.USNM 573677680.
Turtles
Family Geoemydidae
Mauremys reevesii.USNM 573649.
Appendix II. Glossary

Glossary of relevant scientific terms. (doi: 10.3897/zookeys.109.1439.app) File for-
mat: Microsoft Word (doc).
Explanation note: A glossary of relevant scientific terms is provided to readers unfa-

miliar with some of the terms used in our report.
Copyright notice: This dataset is made available under the Open Database License
(http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL)
is a license agreement intended to allow users to freely share, modify, and use this Dataset
while maintaining this same freedom for others, provided that the original source and
author(s) are credited.

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A peer-reviewed open-access journal

ZooKeys 109: 1986 (2011)

The herpetofauna of Timor-Leste: a first report

Corresponding author: Hinrich Kaiser (hinrich.kaiser@vvc.edu)

Materials and methods

Locality District Locality Elevation (m) GPS1

Current Name Synonyms (Authority) Altitude (m) Habitat(s) Frequency

Rana tigerina var. verruculosa (Roux 1911)

Current Name Synonyms (Authority) Altitude (m) Habitat(s) Frequency

Cyrtodactylus sp. Cyrtodactylus sp. (VVC) 600 TEF, CP rare

Eremiascincus antoniorum Glaphyromorphus antoniorum (Greer 1990)

Lygosoma (Omolepida) antoniorum (S)

Current Name Synonyms (Authority) Altitude (m) Habitat(s) Frequency

Elaphe timoriensis (Ba)

Lachesis gramineus (BF98, dR17, Manacas 1972)

Current Name Synonyms (Authority) Altitude (m) Habitat(s) Frequency

fuge tubes containing 95% non-denatured ethanol (EtOH). Ancillary collections of

Frogs and Toads (Order Anura)

Family Dicroglossidae Fork-tongued Frogs

Limnonectes timorensis (Smith, 1927)

Family Hylidae Treefrogs

Family Rhacophoridae Afro-Asian Foam-nest Treefrogs

Lizards (Order Lacertilia)

Collection and natural history. We captured four specimens of D. timoriensis and

given as Draco Timoriensis Pron. However, Pron never published a description of

Family Gekkonidae True Geckos

Gehyra cf. mutilata (Wiegmann, 1834)

Common names. (E) Mutilated Gecko, Stump-toed Gecko, Tender-skinned Gecko.

Gekko gecko (Linnaeus, 1758)

Common names. (E) Tokay Gecko. (T) Toke.

Hemidactylus frenatus Schlegel, 1836

Hemidactylus platyurus (Schneider, 1792)

Family Scincidae Skinks

Cryptoblepharus leschenault (Cocteau, 1832)

Eutropis cf. multifasciata (Kuhl, 1820)

Lamprolepis cf. smaragdina (Lesson, 1826)

f. barbouri, S. f. weberi) additional comparative work is necessary to determine its exact

Family Varanidae Monitor Lizards

Snakes (Order Serpentes)

Aipysurus apraefrontalis*, A. duboisii*, A. foliosquama*, A. fuscus*, A. laevis*, Astrotia stokesii,

Family Colubridae Typical Snakes

Dendrelaphis inornatus timorensis (Smith, 1927)

Lycodon capucinus (Boie, 1827)

Lycodon subcinctus Reinwardt, 1827

Family Homalopsidae Oriental and Australasian Mudsnakes

as C. cf. rynchops. The genus Cerberus is currently undergoing a taxonomic evaluation

Family Typhlopidae Blindsnakes

Family Viperidae True Vipers and Pitvipers

Identification. This snake can be distinguished by both morphology and typical

Turtles (Order Testudines)

Family Chelidae South American and Australasian Side-necked Turtles

Family Geoemydidae South American and Asian Pond Turtles

Crocodiles (Order Crocodilia)

Crocodylus porosus Schneider, 1801

Common names. (E) Saltwater Crocodile, Indo-Pacific Crocodile, Estuarine Croco-

when outlining pressures on the environment in tropical developing countries (natural

Helfenberger N (2001) Phylogeneric relationships of Old World ratsnakes based on visceral

Schlegel H (183744) Abbildungen Neuer oder Unvollstndig Bekannter Amphibien, Nach

Appendix I. Specimens Examined

Frogs and Toads

Appendix II. Glossary

Explanation note: A glossary of relevant scientific terms is provided to readers unfa-

Vous aimerez peut-être aussi

Aipysurus apraefrontalis, A. duboisii, A. foliosquama, A. fuscus, A. laevis*, Astrotia stokesii,